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BMY COTAXON
THE INTERNATIONAL JOURNAL OF FUNGAL TAXONOMY & NOMENCLATURE
VOLUME 105 JULY-SEPTEMBER 2008
Peter Roberts
Heterochaete tenuicula
sp. nov. ( p. 144, Fig 4.)
ISSN 0093-4666 MYXNAE 105: 1-500 (2008)
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MYCOTAXON
Volume 105, pp. 1-5 July-September 2008
A new Pseudocercospora species on Passiflora setacea
ALEXEI C. DIANESE’, ANA M. COSTA’ & JOSE C. DIANESE?™
' alexei.dianese@cpac.embrapa.br
'Embrapa Cerrados, Br-020, Km 18, 73310-970 Planaltina, D.E, Brasil.
“Departamento de Fitopatologia, Universidade de Brasilia
70.910-900 Brasilia, D.F, Brasil.
Abstract — A new Pseudocercospora species, P. passiflorae-setaceae, was detected
causing severe leaf spotting on Passiflora setacea in Brasilia, Distrito Federal, Brazil.
The new species is described, and differences from other Pseudocercospora species on
Passiflora are discussed.
Key words — cercosporoid fungi, Mycosphaerellaceae, tropical fruits, wild passion-
fruit
Introduction
Native Passiflora L. species are common in the Cerrado as well as in other
Brazilian Biomes (Cervi 1997, Nunes & Queiroz 2001, Vieira & Carneiro
2004). Among them P setacea DC. deserves attention being now domesticated
for fruit production. Mendes et al. (1999) listed fungal species associated with
Passiflora in Brazil, but among them species of Pseudocercospora Speg. were
not mentioned. Crous & Braun (2003) revised the names of hyphomycetes
previously referred to as Cercospora Fresen. and Passalora Fr. and published
numerous new combinations with updated descriptions of known cercosporoid
fungi including those on Passiflora species |viz., two Passalora species (Passalora
biformis (Peck) U. Braun & Crous and P. fuscovirens (Sacc.) U. Braun & Crous),
four Cercospora species (C. granadillae Chupp, and three belonging to C. apii
Fresen. s. lat. (viz., C. passifloricola Chupp, C. regalis Tharp, C. truncatella
G.E. Atk.), and three Pseudocercospora species (P. calospilea (Syd.) Deighton,
P. passiflorae U. Braun & Crous, and P. stahlii (F. Stevens) Deighton)] (Chupp
1954, Deighton 1976, Ellis 1976, Crous & Braun 2003). However, the fungus
on P. setacea is quite distinct from the Cercospora and Passalora species
mentioned, which possess conidiogenous cells and conidia conspicuously
' Corresponding author: jcarmine@unb.br
2 ... Dianese, Costa & Dianese
cicatrized. Furthermore, the new species does not coincide with the known
Pseudocercospora species occurring on hosts of the Passifloraceae (Chupp 1954,
Deighton 1976, Braun et al. 1999, Crous & Braun 2003, Farr et al. 2008). The
specimen studied revealed unique features sufficient to be described as a new
Pseudocercospora species.
Materials and methods
Leaves of P. setacea showing strong yellow leaf spotting were collected from a
passion-fruit nursery at Embrapa Cerrados in Brasilia. The sample was dried,
numbered and deposited in the Mycological Collection of the Herbarium of the
University of Brasilia (UB- Col. Micol.). Studies under the stereomicroscope
were followed by observations of squash preparations and sections made with
a freezing microtome. The morphological features were described, measured,
and documented using a Leica DM 2500 microscope coupled with a Leica DFC
490 digital camera connected to a microcomputer. Image capture, editing, and
measurements were made with the help of Leica QWin V3 software. In some
cases, the samples were stained with lacto-glycerol cotton blue and the slides
sealed with nail polish, but most of the photographic work was done without
staining using Nomarski optics. A minimum of fifty replicates of spore and
hyphal measurements were made.
Taxonomy
Pseudocercospora passiflorae-setaceae A.C. Dianese, A.M. Costa & Dianese,
sp. nov. Figs. 1-11.
MycoBank MB 511598
Pseudocercosporae passiflorae similis, sed stromatibus majoribus, (49-) 60-166 (-195)
um diam., hyphis superficialibus cum conidiophoris solitariis evolutis et conidiis longioribus
et latioribus, ad 129 x 5.5 um, ad 15-septatis.
Lesions adaxial, widespread, formed as irregularly scattered yellow solitary leaf
spots, later coalescing into light brown necrotic areas with central black dots;
grayish irregular downy spots on the abaxial surface. IMMERSED MYCELIUM
septate, light brown, within the mesophyll, forming epiphyllous stromata and
hypophyllous superficial mycelium. STROMATA (49-)60-(102)-165(-195)
um diam, epiphyllous, subglobose to globose, brown to dark brown,
subepidermal, erumpent, textura angularis. SUPERFICIAL MYCELIUM exclusively
hypophyllous, strongly branched, intricate, abundant, light grayish brown to
brown, giving rise to conidiophores and conidiogenous cells; hyphae 4-6 um
diam, subhyaline to light bown, septate, smooth, thin-walled. EprpHyLLous
CONIDIOPHORES 7-(14)-21(-31)x2.5-(4)-5 um, numerous, densely fasciculate,
Pseudocercospora passiflorae-setaceae sp. nov. (Brazil) ... 3
Figs. 1-4. Pseudocercospora passiflorae-setaceae on leaves of Passiflora setacea. 1. Yellow irregular
leaf spots on the adaxial face. 2. Gray irregular leaf spots on the abaxial face. 3. Adaxial lesion
showing small dots (stromata) in the central portion of the adaxial side. 4. Abaxial lesions covered
by velvety gray superficial mycelium.
formed on stromata, geniculate, light brown, septate, smooth, HyPOPHYLLOUS
CONIDIOPHORES Solitary, arising from superficial hyphae, lateral or terminal,
aseptate, i.e. conidiophores reduced to conidiogenous cells. CONIDIOGENOUS
CELLS integrated, terminal, 20-50 um long, sympodial; conidiogenous loci
flattened, inconspicuous. Conip1a solitary, cylindrical, rarely obclavate-
cylindrical, 43-(86)-129 x 3-(3.5)-5.5 um, 5-15-septate, solitary, sub-hyaline
to light brown, thin-walled, smooth; apex subacute to subobtuse; base truncate
to somewhat obconically truncate, 3-4.5 um wide; hila neither thickened nor
darkened.
SPECIMEN EXAMINED. BRAZIL. DISTRITO FEDERAL: Planaltina. CENTRO DE PESQUISA
AGROPECUARIA DO CERRADO (CPAC), Km 18, BR-20 HiGHWay NorTH, on living leaves
of Passiflora setacea, 12 Feb 2008, leg. Alexei de Campos Dianese 95, holotype. (UB- Col.
Micol. 20872).
4 ... Dianese, Costa & Dianese
Figs. 5-6. Pseudocercospora passiflorae-setaceae on leaves of Passiflora setacea. Cross section of
stromata with conidiophores on infected leaves showing the trans-mesophyllic mycelium (black
arrows), and also the emerging hypophyllous superficial mycelium (white arrow).
Figs. 7-10. Pseudocercospora passiflorae-setaceae. 7. Hypophyllous intricate superficial mycelium.
8. Branching of the superficial mycelium with terminal conidiogenous cells. 9. Superficial
mycelium (left arrow) bearing a geniculate conidiophore (right arrow) with a 10-septate conidium.
10. Conidia.
Discussion
All three known Pseudocercospora species on Passiflora are easily segregated
from the new species, first of all, because they all lack an external hypophyllous
mycelium with solitary conidiophores. Due to well-developed epiphyllous
stromata with fasciculate conidiophores and similar conidia, Pseudocercospora
passiflorae resembles P._ passiflorae-setaceae, but the stromata are much
smaller, and the conidia are shorter and narrower, usually 30-80 x 2-4
um, only with up to 10 septa (Crous & Braun 2003). The second species on
Passiflora, Pseudocercospora stahlii, was originally described from Puerto Rico
as Helminthosporium stahlii F. Stevens in 1917, and later found in the Virgin
Islands and Dominican Republic. This species is also known from 16 countries
in Asia and Australia mainly affecting Passiflora foetida (Hsieh & Goh 1990,
Hyde & Alcorn 1993, Liu 1977, Guo t al. 1998, Shaw 1984, Thaung 1984,
Zhuang 2001). Pseudocercospora stahlii differs from P. passiflorae-setaceae by
having amphigenous fasciculate conidiophores, smaller stromata (20-40 um),
and short, broad conidia with up to seven septa (Deighton 1976). Finally,
Pseudocercospora passiflorae-setaceae sp. nov. (Brazil) ... 5
P. calospilea is clearly different from the new species by its smaller hypophyllous
stromata (25-35 um diam.), hyaline to subhyaline, cylindrical-obclavate, 1-7-
septate, smaller conidia (20-60 x 2-4 um). Based on the clear morphological
differences, the introduction of a new species for the Pseudocercospora on
Passiflora setacea is justified.
Acknowledgements
The authors thank Embrapa, CNPq and Fundacao Banco do Brasil for financial support,
and Prof. Mariza Sanchez for assistance with the herbarium work, and C.A. Inacio for
help with the photographic work. Also sincere thanks are given to Drs. Uwe Braun and
Roland Kirschner for the pre-submission reviews of our manuscript.
References
Cervi AC. 1997. Passifloraceae do Brasil. Estudo do género Passiflora L., Subgénero Passiflora.
Fontqueria 14:1-92.
Chupp C. 1954. Monograph of the fungus genus Cercospora. Published by the Author: Ithaca, New
York (USA). 667 pp.
Crous PW, Braun U. 2003. Mycosphaerella and its anamorphs - 1. Names published in Cercospora
and Passarola: Centraalbureau voor Schimmelcultures: Ultrecht (The Nederlands). 571 pp.
Deighton FC. 1976. Studies on Cercospora and allied genera. VI. Pseudocercospora Speg., Pantospora
Cif. and Cercoseptoria Petr. Mycological Papers 140:1-168.
Ellis MB. 1976. More dematiaceous hyphomycetes. Commonwealth Mycological Institute: Kew,
Surrey (England). 507 pp.
Farr DE, Rossman AY, Palm ME, McCray EB. 2008. Fungal Databases, Systematic Mycology and
Microbiology Laboratory, ARS, USDA. Retrieved February 14, 2008, from http://nt.ars-grin.
gov/fungaldatabases/.
Guo Y-L, Liu X-J, Hsieh W-H. 1998. Pseudocercospora. Flora Fungorum Sinicorum. Vol. 9. Science
Press: Beijing (China). 474 pp.
Hsieh W-H, Goh T-K. 1990. Cercospora and Similar Fungi from Taiwan. Maw Chang Book Co.:
Taipei (Taiwan). 376 pp.
Hyde KD, Alcorn JL. 1993. Some disease-associated microorganisms on plants of Cape York
Peninsula and Torres Strait Islands. Australasian Plant Pathology 22: 73-83.
Liu PSW. 1977. A supplement to a host list of plant diseases in Sabah, Malaysia. Phytopathological
Papers 21: 1-49.
Mendes MAS, Silva VL, Dianese JC, Ferreira MASV, Santos CEN, Neto EG, Urben AEF, Castro C.
1998. Fungos em Plantas no Brasil. Embrapa - SPI / Embrapa - Cenargen: Brasilia (Brazil).
569 pp.
Nunes TS, Queiroz LP. 2001. A Familia Passifloraceae na Chapada Diamantina, Bahia, Brasil.
Sitientibus Série Ciéncias Bioldgicas 1: 33-46.
Shaw DE. 1984. Microorganisms in Papua New Guinea. Department of Primary Industry Research.
Bulletin 33: 1-344.
Thaung MM. 1984. Some fungi of Cercospora complex from Burma. Mycotaxon 19: 425-452.
Vieira MLC, Carneiro MC. 2004. Passiflora spp. Passionfruit. In: Litz, R. Biotechnology of fruit and
nut crops. Oxford, CABI: Wallingford. p. 436-453.
Zhuang W-Y. 2001. Higher Fungi of Tropical China. Mycotaxon Ltd., Ithaca, NY. 485 pp.
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Volume 105, pp. 7-10 July-September 2008
Type studies on South American Strophariaceae:
2. Pholiota trinitensis is transferred to Stropharia
VAGNER G. CORTEZ
cortezvg@yahoo.com.br
Universidade Federal do Rio Grande do Sul
Programa de Pés-graduacdo em Botanica
Av. Bento Goncalves, 9500, 91501-970, Porto Alegre, RS, BRAZIL
Abstract — The holotype of Pholiota trinitensis (= Hypholoma trinitensis) was
examined, and the presence of numerous acanthocytes in its rhizomorphs necessitates
its transfer to the genus Stropharia. The new combination, S. trinitensis, is proposed.
The taxonomic discussion is accompanied by a brief description and illustrations of
relevant microscopic features.
Key words — Agaricales, Naematoloma, neotropical mycobiota, nomenclature
This new contribution to type studies on South American Strophariaceae Singer
& A.H. Sm. (Cortez 2008) presents results from the type study of Pholiota
trinitensis. Although originally described from Central America (Trinidad
Isl].), there are known records for the species from Brazil (Pegler 1997 from
Sao Paulo State; de Meijer 2006 from Parana State; Bononi et al. 2007 from
Mato Grosso do Sul State) and Argentina (Lechner et al. 2006). This species was
described initially in Pholiota (Fr.) P. Kumm., later transferred to Hypholoma
(Fr.) P. Kumm. (Pegler 1983), and then to Naematoloma P. Karst. (Singer 1986).
However, the characters listed by Dennis and Pegler suggest that the species
belongs to Stropharia (Fr.) Quél. following the current concept proposed
by Cortez & Silveira (2008). The author therefore considered it necessary to
examine the holotype to clarify its generic position.
The type specimens received from Kew were in fragments, consisting of
pieces of two or three basidiomes. Abundant white rhizomorphs were easily seen
on first observation. Pegler (1983), who studied both type and fresh specimens,
provided a detailed macroscopic description that noted a 30-75 mm, convex
to umbonate, cinnamon-brown to ochraceous-buff, hygrophanous pileus
with white velar remnants appendiculate on the margin; adnate, pale grey to
tobacco-brown, moderately crowded lamellae; a 30-50 x 5-8 mm, cylindrical,
ochraceous cinnamon, striate stipe with abundant white rhizomorphs at the
8 ... Cortez
base; a cream colored, fibrillose to floccose veil forming an indistinct annular
zone on upper stipe; and a dark maroon.
Prior to examination, thin sections of dried basidiomes from the holotype
were mounted in 5% KOH and 1% Congo red solutions. The following
characters were observed:
BASIDIOSPORES 6-7.5 X 3.7—4.6(—5) wm, Q= 1.45-1.78, Q = 1.65, n= 42, elliptic
to phaseoliform in side view, ovoid to elliptic in frontal view, with a smooth
and thick wall, and a reduced germ-pore (Fig. 4); BAsIDIA 17-21 x 6-7.5 um,
subclavate, with a medial constriction, 4-spored (Fig. 3); PLEUROCYSTIDIA 38-59
x 10-14.2 wm, as chrysocystidia, lageniform to ventricose, with rounded
apex, not mucronate, thin-walled, with greenish amorphous contents (Fig. 2);
CHEILOCYSTIDIA not found, probably collapsed - according to Pegler (1983)
they are 22-27 x 10-17 um, pyriform to globose, hyaline, and thin-walled,
not of the chrysocystidia type; PILEIPELLIS composed of hyaline, smooth and
thin-walled hyphae, 6-9.5 um diam. on a hypodermium formed by brownish,
inflated hyphae, 14-25 wm diam. (Fig. 1); HYMENOPHORAL TRAMA subregular,
composed by yellowish interwoven hyphae; CAULOCYSTIDIA 27-47 x 9-12
um, as chrysocystidia, scattered or in fascicles on stipe apex, and similar to the
pleurocystidia; ACANTHOCYTES very abundant on rhizomorphs surface (Fig.
5):
The combination of the above characters makes generic assignment in
Strophariaceae difficult, for the spore print suggests Pholiota while the subcellular
hypodermium is more closely related to Hypholoma (= Naematoloma).
However, the presence of acanthocytes easily supports placement of the
species in Stropharia. This morphological feature, which is unique to the genus
(Farr 1980), has been recently useful in diagnosing other Stropharia species
(Cortez 2008; Cortez & Silveira 2007, 2008; Desjardin & Hemmes 2001;
Norvell & Redhead 2000). Recent research (Luo et al. 2006) suggests that in
S. rugosoannulata Farl. ex Murrill acanthocytes function as a nematode-
attacking device unique to the genus.
Pholiota trinitensis must be compared with other neotropical members of
Stropharia previously classified in Pholiota section Albivelatae A.H. Sm. &
Hesler, including S. apiahyna (Speg.) Cortez & R.M. Silveira and S. varzeae
(Singer) Cortez. Stropharia apiahyna differs in producing a smaller pileus
(< 30 mm), a membranous annulus, and longer cheilocystidia (Cortez & Silveira
2008). Stropharia varzeae, which also has membranous annulus, produces
cheilocystidia as chrysocystidia (Cortez 2008). Stropharia acanthocystis Cortez
& R.M. Silveira, from Araucaria angustifolia (Bertol.) Kuntze forests in south
Brazil, is similar in basidiospore size and cheilocystidial shape. However, in
S. acanthocystis the cheilocystidia have thickened walls, pleurocystidia are
Stropharia trinitensis comb. nov. (Brazil) ... 9
Figs. 1-5. Stropharia trinitensis - Holotype: K(M) 147682.
1. Pileipellis and hypodermium. 2. Pleurocystidia. 3. Basidia. 4. Basidiospores 5. Acanthocytes.
clavate, caulocystidia are of three types (leptocystidia, chrysocystidia, and
acanthocytes), and (unique within Stropharia) acanthocytes are also present as
hymenial cystidia (Cortez & Silveira 2007).
Naematoloma amazonicum Singer, described by Singer (1973) from
Brazil, was considered by Pegler (1983, as Hypholoma) and Singer (1986,
as Naematoloma) as a synonym of P. trinitensis. This synonymy only will be
confirmed after the study of the holotype of N. amazonicum.
Based on all above cited characters, P. trinitensis is considered a good species
of Stropharia and the new combination is introduced:
10 ... Cortez
Stropharia trinitensis (Dennis) Cortez, comb. nov.
MycoBank MB 511476
BasionyM: Pholiota trinitensis Dennis, Kew Bull. Add. Ser. 3: 467, 1970.
SynonyMs: Hypholoma trinitense (Dennis) Pegler, Kew Bull. Add. Ser. 9: 509, 1983.
Naematoloma trinitense (Dennis) Singer, Agar. Mod. Taxon.: 564, 1986.
EXAMINED SPECIMEN: TRINIDAD ISL. St. Augustine, 27.IX.1949, R.W.G. Dennis 78 —- K
(M) 147682 (holotype).
Acknowledgements
The author is grateful to the herbarium Kew for loan of the type collection. Special
thanks to Dr. Maria A. Neves (Universidade Estadual de Feira de Santana, Brazil) and
Dr. Lorelei L. Norvell (Pacific Northwest Mycology Service, USA) for critical review of
the manuscript. CNPq (Brazil) is acknowledged for financial support.
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do Pantanal de Rio Negro, MS. Pp. 251-253. In: Barbosa LM, Santos Jr. NA (eds.). A Botanica
no Brasil: pesquisa, ensino e politicas publicas ambientais. Sao Paulo (Brazil): Instituto de
Botanica, SEMA-SP.
Cortez VG. 2008. Type studies on South American Strophariaceae. 1. Pholiota varzeae, from
Brazilian Amazon. Mycotaxon 103: 137-140.
Cortez VG, Silveira RMB. 2007. A new species of Stropharia with hymenial acanthocytes. Mycologia
99: 13 5= 158.
Cortez VG, Silveira RMB. 2008. The agaric genus Stropharia (Strophariaceae) from Rio Grande do
Sul State, Brazil. Fungal Diversity 32: 31-57.
de Meijer AAR. 2006. Preliminary list of the macromycetes from the Brazilian State of Parana. Bol.
Mus. Bot. Mun., Curitiba 68: 1-55.
Dennis RWG. 1970. Fungus Flora of Venezuela and adjacent countries. Kew Bull. Add. Ser. 3:
125318
Desjardin DE, Hemmes DE. 2001. Agaricales of the Hawaiian Islands - 7. Notes on Volvariella,
Mycena Sect. Radiatae, Physalacria, Porpoloma and Stropharia. Harv. Pap. Bot. 6: 85-103.
Farr DE 1980. The acanthocyte, a unique cell type in Stropharia (Agaricales). Mycotaxon 11:
241-249.
Lechner BE, Wright JE, Popoff O. 2006. New taxa and new records for Argentina of fungi from
Iguazu National Park, Misiones. Fungal Div. 21: 131-139.
Luo H, Li X, Li G, Pan Y, Zhang K. 2006. Acanthocytes of Stropharia rugosoannulata function as a
nematode-attacking device. Appl. Environ. Microbiol. 72: 2982-2987.
Maerz A, Paul MR. 1930. A Dictionary of Color. New York (USA): McGraw-Hill, 207pp.
Norvell LL, Redhead SA. 2000. Stropharia albivelata and its basionym Pholiota albivelata.
Mycotaxon 76: 315-320.
Pegler DN. 1983. Agaric Flora of the Lesser Antilles. Kew Bull. Add. Ser. 9: 1-668.
Pegler DN. 1997. The Agarics of Sao Paulo, Brazil. London (UK): HMSO/Royal Botanic Gardens,
Kew, 68pp.
Singer R. 1973. Diagnoses Fungorum Novorum Agaricalium HI. Beih. Sydowia 7: 1-106.
Singer R. 1986. The Agaricales in Modern Taxonomy. 4"° ed. Koenigstein (Germany): Koeltz,
981pp.
MYCOTAXON
Volume 105, pp. 11-18 July-September 2008
Ambispora brasiliensis, anew ornamented species in the
arbuscular mycorrhiza-forming Glomeromycetes
BRUNO TOMIO GOTO!, LEONOR CosTA MAIA! & FRITZ OEHL?
™brunogoto@hotmail.com
‘Departamento de Micologia, CCB, Universidade Federal de Pernambuco
Av. Prof. Nelson Chaves s/n, Cidade Universitaria, 50670-420, Recife, PE, Brazil
*Zurich-Basel Plant Science Center, Institute of Botany, University of Basel
Hebelstrasse 1, 4056 Basel, Switzerland
Abstract — A new dimorphic fungal species in the arbuscular mycorrhiza-forming
Glomeromycetes was found in the Cerrado biome in Minas Gerais (Southeast Brazil)
and is here described from acaulosporoid glomerospores under the epithet Ambispora
brasiliensis. Glomoid glomerospores of this species were found only once, connected
to a germinating acaulosporoid glomerospore. The acaulosporoid glomerospores
are formed on a pedicel branching from the neck of a sporiferous saccule. They are
hyaline to pale yellow in color, globose, 62-95 um in diameter, to subglobose to oval,
59-88 x 69-100(-118) um, and with three spore walls. Crowded irregular pustules,
2.4-7.0(-10.0) um high and 4.2—9.8(—15.0) um wide, are formed on the surface of the
outer wall. The glomoid glomerospores are hyaline to subhyaline, globose, 25-30 um in
diameter, and have a bi-layered spore wall without ornamentation. The acaulosporoid
glomerospores of the new fungus can easily be distinguished from all other species in
the genus Ambispora by spore size and by the distinct pustulate ornamentation.
Key words — Glomeromycota, Ambisporaceae, bimorphic, germination
Introduction
Species of the genus Ambispora C. Walker et al. of the arbuscular mycorrhiza
(AM) forming Ambisporaceae C. Walker et al. (Glomeromycetes) can form
acaulosporoid and glomoid glomerospores (Spain et al. 2006, Walker et al. 2007a,
Walker 2008). For some species only the acaulosporoid morph is known (e.g.
A. jimgerdemannii (N.C. Schenck & T.H. Nicolson) C. Walker), for others only the
glomoid morph (e.g. A. leptoticha (N.C. Schenck & G.S. Sm.) C. Walker et al. and
A. callosa (Sieverd.) C. Walker et al.), while at least A. appendicula (Spain et
al.) C. Walker, A. gerdemannii (S.L. Rose et al.) C. Walker et al. and A. fennica
C. Walker et al. definitely form both morphs (Spain et al. 2006, Walker et
al. 2007a, Walker 2008). There has been some confusion regarding the valid
taxonomic nomenclature of Ambisporaceae and its type genus Ambispora in the
12 ... Goto, Maia & Oehl
recent literature resulting from past incorrect re-combinations and synonymies
(see review by Spain et al. 2006), or obviously illegitimate names attributed by
Spain et al. (2006) at the genus level (see Walker et al. 2007a,b, 2008).
Acaulosporoid glomerospores of Ambispora can quite easily be recognized by
their morphology (Spain et al. 2006) whereas the glomoid morph might be
mistaken with glomerospores of the Glomeraceae. Indeed, Walker et al. (2007a)
identified Glomus callosum Sieverd. as belonging to Ambispora only using
molecular biological techniques.
Acaulosporoid glomerospores of an undescribed Ambispora species were
detected in rhizospheric soil from a natural ecosystem of the Brazilian Cerrado
biome in Serra do Cipé (Minas Gerais State, Southeast Brazil). The species,
characterized by a distinct pustulate ornamentation on the outer spore wall, is
presented here under the epithet Ambispora brasiliensis.
Material and methods
Soil sampling and culturing of AM fungi
Soils were sampled in the Cerrado biome of Serra do Cip6o (Minas Gerais
State, Brazil) from the rhizosphere of typical native plant vegetation, mainly
consisting of Vellozia caruncularis Mart. ex Seub., in July 2004. The site is
located at 19° 06'S and 43° 35°W. Soil pH (H,O) was 4.7.
For culturing the native AM fungal communities, 50 g of the field soil were
mixed with 400 g autoclaved sand-vermiculite substrate (1:1 w/w, pH 5.5) and
filled into 500 ml pots. Sorghum bicolor (L.) Ménch was used as bait plant.
The culture pots were maintained at the greenhouse of the Department of
Mycology, Universidade Federal de Pernambuco, Recife. Additionally, multiple
glomerospores of the new species were separated and used as infective propagules
in single species cultures using the same pot size and sand-vermiculite culture
substrate as described above. So far, all trials failed to establish a successful
symbiosis with the new AMF species both in bait cultures and in single species
cultures.
Morphological analyses
Acaulosporoid glomerospores were extracted from field soils and the
culture substrates by wet sieving (Gerdemann & Nicolson 1963) and sucrose
centrifugation (Jenkins 1964). The spores were thereafter isolated under a
stereo microscope and mounted in polyvinyl-lacto-glycerol (PVLG), PVLG
+ Melzer’s reagent (Brundrett et al. 1994) and in water, respectively. For the
description of the glomerospores and the spore walls, the terminology of Spain
et al. (2006) recently presented for species of the genus was used. We use the
term glomerospore for all spores formed by species of the Glomeromycetes, as
proposed by Goto & Maia (2006).
Ambispora brasiliensis sp. nov. (Brazil) ... 13
Description of the new species
Ambispora brasiliensis B.T. Goto, L.C. Maia & Oehl, sp. nov. Figs. 1-9
MycoBank MB 511612
Sporocarpia ignota. Sporae dimorphae; sporae acaulosporoideae hyalinae vel subhyalinae,
globosae (62-95 um in diametro) vel subglobosae vel ovales (59-88 x 69-100(-118)) um,
cum ternibus tunicis stratis pluribus, in solo singulatim formatae appendice lateraliter ex
hypha terminata in sacculo sporangifero; sporae cum tunica exterior, tunica media et tunica
interior: tunica exterior tunica mediaque coniuncta sacculi et hyphae; tunica exterior
stratis tribus in totum 2.7-3.8 um crassa, stratum exterius tunicae exterioris hyalinum,
evanescens, 0.4-0.7 um crassum; stratum secundum hyalinum at flavo-pallidum, 2.2-3.2
um crassum, pustulibus irregularibus ornatum altis 2.4—7.0(-10)um et 4.2-9.8(-15) um
latis ornatum; stratum interius tunicae exterioris hyalinum at flavor-pallidum, subtile;
tunica media duobus stratis hyalinis in totum 2.8-5.2 um crassa; tunica interior de novo
formans stratis duabus vel tribus, in totum 2.2-2.9 um crassa. Sporae glomoidae hyalinae
vel subhyalinae, 25-30 um in diameterum, stratis duobus. Holotypus hic depositus 78879:
URM.
EryMo.ocy: brasiliensis referring to Brazil, the country where the species was found.
GLOMEROSPORE FORMATION. Sporocarp formation is unknown. The species
is dimorphic producing acaulosporoid and glomoid morphs. Acaulosporoid
glomerospores form singly in soils upon a short pedicel arising laterally on the
tapering hyphal neck of a sporiferous saccule (Figs. 1-3). The acaulosporoid
glomerospores form three spore walls: an outer wall (ow), a middle wall (mw)
and an inner wall (1w) (Figs. 4-9). Glomoid glomerospore formation was
observed once on a hypha connected to a germinating acaulosporoid morph.
SPORIFEROUS SACCULE OF ACAULOSPOROID GLOMEROSPORESare hyaline to light
yellow, globose to subglobose, 65-115 um in diameter and formed terminally
on a hypha (Figs. 1-2). The saccule wall generally is bi-layered with a hyaline
to light yellow, rapidly degrading, evanescent outer layer (0.5-0.9 ttm thick)
and a hyaline to subhyaline, semi-persistent inner layer (0.6-1.5 um thick).
The hyphal neck of the sporiferous saccule is 10-17 um wide at the saccule
base tapering to 8-12 um at the point where the pedicel of the acaulosporoid
glomerospore branches, narrowing further to 5-8 um at a distance of 40-120
um from the pedicel. A septum arises in the tapering hyphal neck distal to the
saccule where the pedicel forms. The sporiferous saccule generally collapses
and detaches from the mature spore while the pedicel often persists on the
glomerospore.
ACAULOSPOROID GLOMEROSPORES form singly on a short hyphal pedicel (Fig.
3). The spores are globose (62-95 um in diameter) to subglobose to oval, 59-88 x
69-100(-118), hyaline to subhyaline in water and generally become light yellow
to ochre-yellow in PVLG-based mountants. The two wall layers of the saccule
continue on the glomerospore differentiating into two separate glomerospore
walls: a two to three layered outer wall (ow) and a bi-layered middle wall (mw).
14 ... Goto, Maia & Oehl
A two-to-three layered inner wall (tw) forms de novo, presumably after the
spore pore has been closed by the inner layer of the middle wall (mw12) which
differentiates from MwL1. The rw functions as a germinal wall.
OUTER WALL generally consists of three layers (Figs. 4-8). The outer layer (OWL1)
is hyaline to subhyaline, evanescent, 0.4-0.7 um thick, and often is difficult to
detect, as it tightly adheres to owL2 (Fig. 8). OWL2 is hyaline to light yellow and in
total 2.2-3.2 um thick and crowded with conspicuous, convex pustules (Fig. 3-8).
The pustules vary greatly in shape and size and are generally 2.4—7.0(-10)
um wide and 4.2-9.8(-15) high (Figs. 4-7). The inner layer (OWL3) is thin
(0.3-0.6 um), hyaline to subhyaline and tightly attached to owL2 making it
difficult to detect. In older spores, the outer wall might degrade completely, but
usually some fragments remain on the surface of the middle wall.
MIDDLE WALL is bi-layered and in total 2.8-5.2 um thick (Figs. 4-6; Figs. 8-9).
MWLI is hyaline, 1.3-2.6 um thick and continuous with the persistent inner
layer of the pedicel. A septum arising in the pedicel from MWwL1 during spore
formation was rarely detected. MWL2 is also hyaline, 1.5-2.6 tim thick, tightly
adherent to MwL1 and generally closes the pore of the pedicel.
INNER WALL is hyaline and three-layered but generally only the central layer
(IWL2; Figs. 4-6; Fig. 8—9) or the inner two layers (IwL2 and IWL3) are visible. The
outer layer IWL1 is 0.3-0.6 um and difficult to detect, as it is tightly adherent to
the central layer IwL2 which is 1.5-2.2 um thick. The innermost layer is 0.4—0.8
tum thick and also tightly adherent to rwL2 but sometimes separates slightly
showing several thin folds when pressure is applied.
PEDICEL OF ACAULOSPOROID GLOMEROSPORE branches from the hyphal neck
of the sporiferous saccule in about 40-130 um distance from the globose
saccule terminus. The pedicel is (5—)7—16(-25) um long, and 8-13 um wide at
the glomerospore base tapering to 5-9(-13) um at the connection to the hypha.
The two wall layers of the hyphal neck of the sporiferous saccule differentiate
into the outer and the middle wall of the pedicel (ow and Mw). On mature
spores, the pedicel often persists resembling a subtending hypha of glomoid
glomerospores (Fig. 5). Sometimes only the ow (Fig. 5) or the Mw (Fig. 6) of
the pedicel persist on the spore; rarely both pedicel walls are found on mature
glomerospores. The ow sometimes forms a collar (Fig. 7) on the glomerospore
(8.5-14 um in diameter). The ow, if present on the pedicel, is 1.2-2.5 um thick
at the glomerospore base tapering to 0.5-0.9 um towards the saccule neck. The
Mw is 1.3-2.1 um at the glomerospore base tapering to 0.9-1.5 um.
GLOMOID GLOMEROSPORES were detected only once on a connecting hypha
about 250 um from a germinating acaulosporoid glomerospore. ‘The hyaline
to subhyaline glomoid glomerospores were bi-layered and about 25-30 um
in diameter. The glomerospore wall was 1.9-2.5 um thick in total, and the
Ambispora brasiliensis sp. nov. (Brazil) ... 15
Dee ser
Figs. 1-9: Ambispora brasiliensis forming acaulosporoid glomerospores with three walls (outer wall
ow, middle wall mw and inner wall rw) upon a short pedicel arising laterally on the neck of a
sporiferous saccule. Fig. 1. Acaulosporoid glomerospore and sporiferous saccule (sac) attached.
Fig. 2. Saccule with two layers: evanescent outer layer continuous with ow of the spore and semi-
persistent inner layer continuous with MwL1 of the acaulosporoid spore. Fig. 3. Pedicel (pcL) and
degrading saccule. Figs. 4-6. Irregular pustulate ornamentation (ORN) on middle layer of ow; MW
has two layers (MwLI + Mw12). Fig. 5. Pcl wall continuous with ow. Fig. 6. Pcl wall continuous
with MwLl. Fig. 7. Glomerospore pore and collar formed on ow. Figs. 8-9. Spore wall structure:
ow with three layers (evanescent owLl, pustulate owL2; owL3 hardly detectable), mw bi-layered
(MwLI1 + MWL2); only central layer evident on Iw.
16 ... Goto, Maia & Oehl
subtending hypha was 2.0-3.1 «sm wide at the base of the glomerospore, where
a septum was not visible.
GERMINATION was observed once on an acaulosporoid glomerospore, where
it initiated on the rw below the pedicel collar. Three germ tubes (2.4-4.0 um
wide) emerged from the center of a lobed germination structure, which was
34 x 15 um in diameter and consisted of 6-7 lobes that were each 12-15 um
long and 5-8 um wide. The germ tubes had grown through the pedicel collar
and branched in about 200-250 ttm distance from the glomerospore. However,
it was not clearly discernable if the lobed germination structure was formed
between Iw and Mw or between the mw and the collar formed by the pedicel
on the ow.
MYCORRHIZAL FORMATION — unknown.
DisTRIBUTION — Brazil: so far only found in the Cerrado biome of Serra do
Cipo, Minas Gerais State.
SPECIMENS EXAMINED — BRAZIL. Minas Gerais. Serra do Cipé. HOLOTYPE (54-
5401) and type specimens (54-5402) deposited at URM, Recife, Pernambuco, Brazil
(accession numbers URM 78879 and 78880). Isotype (54-5402) specimens deposited
at OSC, Corvallis, Oregon, USA (accession number OSC# 134501); and 54-5406 & 54-
5407, deposited at Z+ZT, Zurich, Switzerland (accession number ZT MYC 159).
Discussion
The new fungal species, Ambispora brasiliensis, can easily be identified by
its acaulosporoid morph and can be distinguished from all other Ambispora
species forming acaulosporoid morphs by glomerospore size and by the distinct
pustular ornamentation on the acaulosporoid glomerospore walls.
Acaulosporoid glomerospores of other genera in the Glomeromycetes — namely
Acaulospora, Archaeospora and Otospora (Gerdemann & Trappe 1974, Morton &
Redecker 2001, Palenzuela et al. 2008) — cannot be confused with A. brasiliensis.
Spores of Archaeospora and Otospora have two walls (Palenzuela et al. 2008)
contrasting with Acaulospora and Ambispora, which have three spore walls
(Spain et al. 2006, Walker et al. 2007a, Palenzuela et al. 2007). Glomerospores
of Acaulospora are not formed on a pedicel (= appendix) as known for the
acaulosporoid glomerospore formation of Ambispora. Furthermore, in
Acaulospora the middle spore wall forms de novo (e.g. Oehl et al. 2006) and is
not continuous with the wall of the sporiferous saccule as noted for Ambispora
brasiliensis and other Ambispora species (Spain et al. 2006, Walker et al. 2007a).
Finally, only three Acaulospora species are known to form projections on the -
outer spore wall: Ac. spinosa, Ac. tuberculata and Ac. denticulata, which form —
according to their names — either spiny, tuberculate or dentate ornamentations
(Walker & Trappe 1981, Janos & Trappe 1982, Sieverding & Toro 1987) but not
convex, irregular pustules as in A. brasiliensis.
Ambispora brasiliensis sp. nov. (Brazil) ... 17
We found a glomoid morph of this new species connected via hypha to a
germinating acaulosporoid morph of A. brasiliensis only in one occasion. Such
glomoid glomerospore formation during germination of acaulosporoid spores
was also found for A. appendicula (Spain pers.comm. 2008; Goto & al. unpub.
obs.). In addition to the glomoid morph, swollen hyphal tips on mycelia were
also noted for A. appendicula (Schenck et al. 1984).
We were not able to culture A. brasiliensis but hope to conduct genetic analyses
as soon as clean cultures are available. However, in the absence of genetic data,
A. brasiliensis can be identified morphologically by observing its acaulosporoid
glomerospores. As stated in the introduction, the glomoid morph of some of
the Ambispora spp. is still unknown, and, in any case, Ambispora is more easily
diagnosed by observing the acaulosporoid rather than the glomoid morph.
Acknowledgements
The authors acknowledge, in special, Dr. Ewald Sieverding (Institute of Plant Production
and Agroecology in the Tropics, University of Hohenheim, Stuttgart, Germany) and
Joyce L. Spain (Corvallis, Oregon USA), for reviewing the manuscript and making
helpful comments and suggestions, and Dr. Nanuza Luiza de Menezes (Universidade
de Sao Paulo, Brasil) for providing the opportunity to visit the Serra do Cipé, MG.
This work was supported by the Conselho Nacional de Desenvolvimento Cientifico
e Tecnoldgico (CNPq) and Coordenacao de Aperfeigoamento de Pessoal de Nivel
Superior (CAPES) that provided, respectively, a research grant to Leonor C. Maia and a
PhD scholarship to Bruno T. Goto. This work was also supported by the Universidade
Federal de Pernambuco, which provided a grant to F. Oehl as ‘visiting professor’
Literature cited
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Publications, University of Guelph, Guelph, Ontario, Canada.
Gerdemann JW, Nicolson TH. 1963. Spores of mycorrhizal Endogone species extracted from soil by
wet sieving and decanting. Trans. Br. Mycol. Soc. 46: 235-244.
Gerdemann JW, Trappe JM. 1974. The Endogonaceae in the Pacific Northwest. Mycologia Mem.
No. 5. 76 p.
Goto BT, Maia LC. 2006 Glomerospores: a new denomination for the spores of Glomeromycota,
a group molecularly distinct from Zygomycota. Mycotaxon 96: 129-132.
Janos DP, Trappe JM. 1982. Two new Acaulospora species from Tropical America. Mycotaxon 15:
DES-522.
Jenkins WR. 1964. A rapid centrifugal-flotation technique for separating nematodes from soil.
Plant Dis. Rep. 48: 692.
Morton JB, Redecker D. 2001. Two new families of Glomales, Archaeosporaceae and Paraglomaceae,
with two new genera Archaeospora and Paraglomus, based on concordant molecular and
morphological characters. Mycologia 93: 181-195.
Nicolson TH, Schenck NC. 1979. Endogonaceous mycorrhizal endophytes in Florida. Mycologia
71: 178-198.
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Oehl F, Sykorova Z, Redecker D, Wiemken A, Sieverding E. 2006. Acaulospora alpina, a new
arbuscular mycorrhizal fungal species characteristic for high mountainous and alpine
grasslands of the Swiss Alps. Mycologia 98: 286-294.
Palenzuela J, Ferrol N, Boller T, Azcén-Aquilar C, Oehl F. 2008. Otospora bareai, a new fungal
species in the Glomeromycetes from a dolomitic shrub-land in the National Park of Sierra de
Baza (Granada, Spain). Mycologia 100: 282-291.
Schenck NC, Spain J, Sieverding E, Howeler RH. 1984. Several new and unreported vesicular-
arbuscular mycorrhizal fungi (Endogonaceae) from Colombia. Mycologia 76:685-699.
Sieverding E, Toro S. 1987. Acaulospora denticulata sp. nov. and Acaulospora rehmii sp. nov.
(Endogonaceae) with ornamented spore walls. Angew. Bot. 61: 217-223.
Spain JL, Sieverding E, Oehl F. 2006. Appendicispora, a new genus in the arbuscular mycorrhizal-
forming Glomeromycetes, with a discussion of the genus Archaeospora. Mycotaxon 97:
163-182.
Walker C. 2008. Ambispora and Ambisporaceae resurrected. Mycol. Res. 112: 297-298.
Walker C, Trappe JM. 1981. Acaulospora spinosa sp. nov. with a key of Acaulospora. Mycotaxon
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Walker C, Vestberg M, Demircik F, Stockinger H, Saito M, Sawaki H, Nishmura I, Schiissler A. 2007a.
Molecular phylogeny and new taxa in the Archaeosporales (Glomeromycota): Ambispora fennica
gen. sp. nov., Ambisporaceae fam. nov., and emendation of Archaeospora and Archaeosporaceae.
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MYCOTAXON
Volume 105, pp. 19-22 July-September 2008
Lichens of Malatya, Elazig and Adiyaman provinces (Turkey)
MEHMET CANDAN* & AYSEN OZDEMIR TURK
mecandan@anadolu.edu.tr e& aturk@anadolu.edu.tr
Department of Biology, Faculty of Science, Anadolu University
Eskisehir, Turkey
Abstract—The lichenized and lichenicolus fungi of Malatya, Elazig and Adiyaman
provinces of Turkey, based on the authors’ collections and literature records, are listed,
of which 20 are new for Turkey and 187 are new for the study area. As a consequence
of this study, there are 315 infrageneric taxa recorded from these provinces. The full
Checklist can be viewed at http://www.mycotaxon.com/resources/weblist.html
Key words—lichenized fungi, new records, biodiversity
An increasing number of studies on the lichen flora of Turkey have been carried
out in the last two decades, most focused on central, western and northern Turkey
where even today some areas remain unexplored. The present floristic study of the
previously overlooked Malatya, Elazig and Adityaman provinces redresses to some
extent this situation.
The first lichen records of six independent researchers from the study area,
concerned with the distribution and uses of manna lichens undertaken during
1864-1897, are reviewed by Donkin (1981). Steiner (1921) reported 74 taxa from
Elazig and Adiyaman provinces, based on Handell-Mazettis collections from
Mesopotamia, Syria and Iraq. Szatala (1960) recorded 8 lichens from Elazig
Province, as well as records from other parts of Turkey. Szatala’s paper (1960)
based on the lichen collection of zoologist Victor Pietschmann, all contained
lichen records from Syria and Iraq. Recently, Candan & Ozdemir Tiirk (2000)
recorded 50 infrageneric taxa from eastern part of Malatya City. Other records
for the study area are two exsiccates (Lumbsch & Feige 1999, John 1999), two by
Degelius (1954), one by Steiner & Poelt (1982), one by Timdal (1991) and new
lichenicolous fungi by Halici et al. (2007a,b) and Halici & Candan (2007).
As in other parts of Turkey, the natural vegetation of Malatya, Elazig and
Adiyaman provinces has been subjected to anthropogenic stress for many years,
mainly due to heavy grazing, afforestation, farming and urbanisation. Furthermore,
many large dams have been built in southeastern Anatolia during the last 30 years
and others are currently under construction. Hence, a relatively large portion of
the study area is covered by dam lakes. As well as determining the current lichen
20 ... Candan & Tiirk
Figure 1. Map of the study area
diversity of Malatya, Elazig and Adiyaman provinces, this article also includes
previous records from the region dating back to the 1860s. It is also hoped that
this study will provide a database for monitoring the effects of the climate change
on the lichen flora due the development of dams and their resultant lakes.
Stupy AREA: Malatya, Elazig and Adiyaman provinces are located in Anatolia and
are part of the Irano-Turanian Phytogeographical Region of Turkey. According to
Davis (1965), Malatya and Elazig are in the Upper Euphrates and Adiyaman is in
the Mesopotamian area. The study area (Fig. 1) is located between 37°25’ 39°11’
N and 37°25’ 40°21’ E at altitudes ranging from 500 m in the south of Adryaman
Province to 2650 m in the Malatya Mountains lying on the border of Adtyaman
and Malatya provinces. The northern part of the study area, including Malatya
and Elazig provinces, is mainly covered by highlands, the lowest altitudes of c. 800
m being found in valleys.
The study area also covers the eastern side of the Nurhak Mountains, the
Malatya Mountains, the Maden Mountains and Akdag located in the Southeast
Taurus Mountains, the highest altitudes being the summits of Akdag (2650 m),
Bey (2545 m), Hazar (2347 m), Akcababa (2052 m) and Hasan (1864 m). The
south side of Yama Mountain (2402 m), situated between Malatya and Sivas
provinces, is also included in the study area. The Upper Firat, Tohma Stream,
Siirgii Stream, Sultan Creek, Murat, and Dicle Valleys are located between the
high mountains. Although Hazar Lake is the largest natural lake in the area, the
lakes of Keban, Karakaya, Atatiirk, Siirgii and Cat Dams cover a much larger area
collectively.
The rich topographical diversity of the study area provides varied climatic
conditions, which are further modified the effects of the dam lakes; to date, no
Lichens of Malatya, Elazig, & Adiyaman... 21
research has been carried out to determine these effects. The study area has a
Meditteranean climate. According to Emberger’s principles, Akman (1990)
reported that precipitation-temperature coefficients (Q,) are 36.2, 39.6 and 75.4
and aridity indices (S) are 0.7, 0.5 and 0.2 for Malatya, Elazig and Adryaman
provinces, respectively.
The dominant vegetation of the region is steppe in character. For anthropological
reasons, no large forests are to be found in the study area; Quercus spp. are sparsely
distributed. The most eastern distribution of Pinus brutia Ten. in Turkey is
located at the junction of the borders of Kahramanmaras, Malatya and Adiyaman
provinces which coincides with the southeast part of the study area. In spite of
poor forest cover, there is intensive apricot (Prunus armeniaca L.) agriculture
in some parts of Malatya Province and pistacio (Pistacia vera L.) agriculture in
Adtyaman Province. Wild Prunus, Crataegus and Rosa spp. are also common in
the study area, and Salix spp., Populus spp., and Platanus orientalis L. are found in
the wet habitats such as valleys.
MeTHoboLoGy: This study is based on an evaluation of the authors’ lichen
collections made between 2003 and 2005 and on literature sources relating
to Malatya, Elazig and Adiyaman provinces of Turkey. The specimens were
examined with a Leica MZ6 stereomicroscope and an Olympus BX51 microscope
with standard identification methods for lichens and lichenicolous fungi (Purvis
et al. 1992, Wirth 1995). Also TLC analyses of the specimen were carried out
when needed. Voucher specimens are stored in ANES (Herbarium of Anadolu
University, Anadolu University, Faculty of Science, Eskisehir, Turkey).
CONCLUSIONS: Twenty ofthe lichens in the Checklist are new for Turkey: Acarospora
macrospora subsp. murorum, A. peliscypha, Aspicilia emiliae, Buellia imshaugii,
B. pulverulenta, Caloplaca erodens, Dermatocarpon rivulorum, Fulgensia pruinosa,
Immersaria usbekica, Lecanora laatokkaensis, L. reuteri, Placidiopsis custnani,
Placidium tenellum, Polyblastia albida, Rinodina interjecta, R. luridata, R. roscida,
Staurothele bacilligera, Thelidium incavatum, and Xanthoparmelia glabrans.
According to the literature, 128 lichens and lichenicolous fungi have been
previously recorded from the study area. As a consequence of this study, this
number has increased to 315. However, some early papers (Donkin 1981, Steiner
1921, Szatala 1960) provide only vague locality details (many of which have
changed their names), making it difficult to evaluate distributional changes of
lichens.
The study area is a transition region between the Mediterannean and the
Irano-Turanian Phytogeographical Regions, the western and southwestern parts
bordered by the former in Turkey where the distribution of Pinus brutia ends.
This is why some authors delimit this region by the distribution of P. brutia while
others delimit it by the Mediterranean sclerophyll forests, but both have very
similar phytogeographical boundary.
22 ... Candan & Tiirk
Because of the interesting geographical position of the study area, lichens
commonly distributed in Central Asia, Middle East and Europe also occur with
local lichens, leading to a rich lichen diversity. Furthermore the altitude of the
study area starts from 500 m in Adiyaman Province and 800 m in Elazig Province,
and reaches higher than 2000 m in some localities. Therefore, high mountain
lichens also make a significant contribution to the lichen flora of the area. However,
lichen diversity is modified by the limited forest cover which reduces humid and
shade loving species; most lichen epiphytes grow on free-standing trees and are
nitrophilic and heliophilic.
Acknowledgements
We are grateful to Prof. Mark R. D. Seaward for linguistic revision, helpful comments
and reviewing the manuscript, to Dr. P. Divakar for reviewing the manuscript and to
the authors whose names are indicated in the List of Taxa. This study was funded by
Anadolu University Scientific Research Project Unit (grant no. 031043).
Literature cited
Akman Y. 1990. [klim ve Biyoiklim. Ankara, Palme.
Davis P. 1965. Flora of Turkey. Vol: 1. Edinburgh, Edinburgh University Press.
Candan M, Ozdemir Tiirk A. 2000. Orduzu-Malatya bélgesi likenleri. OT Sistematik Botanik
Dergisi 7: 219-230.
Degelius G. 1954. The lichen genus Collema in Europe. Symb. Bot. Upsal. 13 (2): 183, 329.
Donkin RA. 1981. Reports and Comments, The Manna Lichen: Lecanora esculenta. Anthropos 76:
562-572.
Halici1 MG, Candan M. 2007. Notes on some lichenicolous fungi from Turkey. Turkish Journal of
Botany 31: 353-356.
Halici MG, Ozdemir Tiirk A, Candan M. 2007a. New records of pyrenocarpous lichenicolous fungi
from Turkey. Mycotaxon 99: 201-206.
Halici MG, Atienza V, Hawksworth DL. 2007b. Two new Polycoccum (Dothideales, Dacampiaceae)
species from Turkey. Mycotaxon 101: 157-163.
John V. 1999. Lichenes Anatolici Exsiccati. Arnoldia Fasc. 1-3 (No. 1-75): 44.
Lumbsch HT, Feige GB. 1999. Exsiccat. Lecanoroid lichens. Univ. Essen, Essen, Fasc. 5: 1-9.
Purvis OW, Coppins BJ, Hawksworth DL, James PW, Moore DM. 1992. The Lichen Flora of Great
Britain and Ireland. London, Natural History Museum Publications in Association with The
British Lichen Society.
Steiner J. 1921. Lichenes aus Mesopotamien un Kurdistan sowie Syrien und Prinkipo. Gesammelt
von Dr. Heinrich Frh. v. Handell-Mazzetti (wissenschaftliche Ergebnisse der Expedition nach
Mesopotamien 1910). Ann. des Naturhist. Museums in Wien 34: 1-68.
Steiner M, Poelt J. 1982. Caloplaca sect. Xanthoriella, sect. nov: Untersuchungen tiber Xanthoria
lobulata-Gruppe (Lichenes, Teloschistaceae). Plant Systematics and Evolution 140: 151-177.
Szatala O. 1960. Lichenes Turcicae Asiaticae ab Victor Pietscmann collect. Sydowia 14: 312-325.
Timdal E. 1991. 41 monograph of the genus Toninia (Lecideaceae, Ascomycetes). Opera Botanica
110: 1-137.
Wirth V. 1995. Di2 Flechten Baden Wiirttembergs. Teil 1-2, Stuttgart, Ulmer.
MYCOTAXON
Volume 105, pp. 23-27 July-September 2008
Two new anamorphic rust fungi
from northern areas of Pakistan
MUHAMMAD ASIM SULTAN’, IKRAM-UL-HAQ?,
A.N. KHALID? & HAMID MUKHTAR?
asim418@ yahoo.com
‘Department of Biology, University College Lahore, Pakistan
ikrhaq@ yahoo.com
*Department of Botany, Government College University, Lahore, Pakistan
drankhalid@ yahoo.com
*Department of Botany, University of the Punjab, Lahore, Pakistan
hamidwaseer@yahoo.com
*Department of Botany, Government College University, Lahore, Pakistan
Abstract — Two anamorphic rust fungi, Aecidium pakistanicum on Spiraea tomentosa
and Aecidium ikramii on Rosa sp. are described as new. These new discoveries raise the
total of known rust taxa in northern Pakistan to forty-four.
Key words — Basidiomycota, new taxa, Phragmidium tuberculatum
Introduction
Northern areas of Pakistan are bounded by Afghanistan, People’s Republic of
China, Indian-held territory of Kashmir and North West Frontier Province of
Pakistan. The southern slopes near Kashmir lie within the Himalayan mountain
system. Snow covered mountains with deep gorges and narrow valleys surround
the area. Climatic conditions vary widely in the northern areas, ranging from
monsoon influenced moist temperate zone in western Himalayas to arid and
semi-arid cold desert in the northern Karakoram and Hindu Kush. Temperature
in the valley bottoms can vary from extremes of 40°C in summer to less than
10°C in winter (Jacobose 1993).
The rust fungi of Northern areas of Pakistan are poorly known and only forty-
two rust taxa have been published from the region so far (Ahmed 1956a, b;
* Author for correspondence
24 ... Sultan & al.
Gjaerum & Iqbal 1969, Kaneko 1993, Kakishima et al. 1993, Khalid et al. 1995,
Khalid & Iqbal 1996a, b, 1997; Ahmed et al. 1997).
This paper reports two anamorphic rust fungi collected on rosaceous plants,
Spiraea tomentosa L. and Rosa sp., from northern areas of Pakistan during
2001-02
Materials and methods
The host plants, Spiraea tomentosa and Rosa sp. were found infected with rust
fungi. Aecidium pakistanicum on S. tomentosa from Fairy-meadows on July
22, 2001 and Aecidium ikramii on Rosa sp. from Bashu Jungle, Skardu, on
October12, 2002 were collected by the senior author.
Dried herbarium materials were examined under light microscope and
Scanning Electron Microscope (SEM). For LM observations, the hand sections
and spores were mounted in a drop of Lactophenol solution on glass slides,
gently heated to boiling point and then cooled. The measurements of spores
are given in the form: min-max (mean = | standard division). The sections
showing rust stages were observed under a microscope (NIKON YS 100) and
micro photographed by Canon AE-1 programmed camera. For SEM the spores
were attached to specimen stub in a drop of acetone and coated with carbon
with Joel Jee-420 evaporator. The surface structure of spores was observed at 10
kV and photographed with Joel Jsn-6480 LV SEM.
Enumeration of taxa
Aecidium pakistanicum Sultan, I.U. Haq, Khalid & Mukhtar, sp. nov.
MycoBank MB 511527 (Plate 1, Figs A, C)
Pycnia epiphylla, brunneis, dispersa, 0.1-0.2 mm. Aecidia hypophylla, dispersa, aggregate,
revoluto, 0.2-0.7 mm, brunneis, peridio longo, hyalinae, cellula peridii ellipsoideae, 68-
88 x 15-19 um, pariete externo 3.5-5.5 um crasso, verrucoso, pariete interno 5.0-7.5 um
crasso, verrucoso, Aecidiosporae 34-36 x 26-28 um, globosae vel subglobosae, ellipsoideae,
hyalinae, brunneis, verruculoso, Poris obscures. Uredinia et telia ignota.
Holotype: On Spiraea tomentosa L., with 0 & 1 stages, Northern Pakistan, Fairy- Meadows,
July 22, 2001, M.A. Sultan (LAH Herbarium # AS 41, Isotype GCU Herbarium).
EryMo.oey: Based on the name of country Pakistan.
Pycnia epiphyllous, brown, scattered, 0.1-0.2 mm. Aecia hypophyllous,
scattered, sori appear aggregate, in groups, revolute, 0.2-0.7 mm, brown, peridia
long, hyaline. Peridial cells (Fig 1A) ellipsoid, 68-88 x 15-19 um, outer wall 3.5-
5.5 um thick, verrucose, inner wall 5.0-7.5 um thick, verrucose. Aeciospores
(Fig 1B) 34-36 x 26-28 um, globose to subglobose, ellipsoid, hyaline, brown,
densely and finely verruculose, pores obscure. Uredinia and telia unknown.
Aecidium spp. nov. (Pakistan) ... 25
Plate 1. Figs. A-B: Aecidium pakistanicum on Spiraea tomentosa (A) Infected host plant
Bar = 2.0 cm. (B) Aeciospores in Light microscope Bar =160 um. (C) Verruculose aeciospores
in SEM. Bar = 18 um.
COMMENTS — Aecidium pakistanicum is unique among the known species
of Aecidium in having a sac-like sorus and verruculose aeciospores. These
characteristics separate A. pakistanicum from fourteen (Sato & Sato 1984)
described aecium types. A. pakistanicum resembles none of the aecial types
described so far.
Aecidium haqii Sultan, Khalid & Mukhtar, sp. nov. Plate 2, Figs. A-C
MycoBANK MB 511528
Spermogonia, uredinia et telia ignota. Aecidia hypophylla, dispersa, aggregate, caeomoidea,
paraphysibus peripheralis, flavida, 0.1-0.2 mm. Aecidiosporae globosae vel subglobosae,
ellipsoideae, hyalinae, 20-26 x 19-25 um, clavo superne granulis refractivis, parietibus
usque ad lum crassis, poris obscures.
Holotype: On Rosa sp., with I stage, Northern Pakistan, Bashu Jungle near Skardu,
October 12, 2002, M.A. Sultan (LAH Herbarium # AS 39, Isotype GCU Herbarium)
26 ... Sultan & al.
Plate 2. Figs. A-C: Aecidium hagii on Rosa sp. (A) Infected host plant Bar = 2 cm.
(B) Aeciospores in Light microscope Bar = 2 um. (C) Annulate aeciospores in SEM.
Bar = 6.0 um
EtymMo oey: In honor of the renowned botanist Prof. Dr. Ikram-ul-Haq.
Spermogonia, uredinia and telia unknown. Aecia hypophyllous, scattered, in
groups, caeomoid, paraphysis peripheral, yellow, 0.1-0.2 mm. Aeciospores (Fig
2) globose, subglobose, ellipsoid, hyaline, 20-26 x 19-25 um, annulate, wall 1
um thick, pores obscure.
ADDITIONAL SPECIMENS EXAMINED: LAH # 14131, 14132 S. Ahmed, on Rosa webbiana
Wall. ex Royle, Kaghan Valley August 28, 1949.
Aecidium hagii was found on an unidentified Rosa species lacking thorns and
with small leaves. The new species closely resembles Aecidium of Phragmidium
tuberculatum J.H.H. Mill. Both have a caematoid aecidium and similarly
sized aeciospores (see Wilson & Henderson 1966). Both species can be
clearly differentiated on the basis of wall ornamentation of aeciospores. In P
tuberculatum the aeciospores are verrucose and in Aecidium haqii aeciospores
are annulate.
Aecidium spp. nov. (Pakistan) ... 27
Acknowledgements
We sincerely thank Dr. H. B. Gjaerum for help in verification of the taxa. We greatfully
acknowledge Dr. Y. Ono, Ibaraki University, and Dr. S. H. Iqbal, University of Punjab, for
reading the manuscript as pre-reviewers. We extend our thanks to A. R. Niazi for help in
laboratory and to Drs. A. Latif, A. Hameed, M. Ghauri, Government College, University
Lahore, for the SEM photographs. We are grateful to Dr. M. Weif’, Universitat Tubingen,
for help in Latin diagnosis.
References
Ahmed S. 1956a. Uredinales of West Pakistan. Biologia 2(1): 29-101.
Ahmed S. 1956b. Fungi of Pakistan. Biological Society Pakistan Lahore Monograph 1: 1-126.
Ahmed S, Iqbal SH, Khalid AN. 1997. Fungi of Pakistan. Mycological Society of Pakistan,
_ Department of Botany, University of Punjab, Lahore, Pakistan.
Gjaerum HB, Iqbal SH. 1969. Some rust fungi from West Pakistan. Nytt Mag. fiir Botanik 16:
221-223.
Jacobose JP. 1993. Climatic records from Northern Areas of Pakistan. In: Culture Area Karakorum
Newsletter, Tubingen, Germany, 3: 13-17.
Kaneko S. 1993. Parasitic fungi on woody plants from Pakistan. Cryptogamic Flora of Pakistan 2:
149-168.
Kakishima M, Okaen I, Ono Y. 1993. Rust Fungi (Uredinales) of Pakistan collected in 1991.
Cryptogamic Flora of Pakistan 2: 169-179.
Khalid AN, Iqbal SH, Masood A. 1995. New records of Uredinales from Pakistan. Science
International (Lahore) 7(4): 531-532.
Khalid AN, Iqbal SH. 1996a. Additions to the Rust Flora of Pakistan. Pak. J. Bot. 28(1): 114-117.
Khalid AN, Iqbal SH. 1996b. New Rusts from Pakistan. Can. J. Bot. 74: 506-508.
Khalid AN, Iqbal SH. 1997. Puccinia deosaiensis sp.nov on Epipactis helleborine (L.) Crantz. Can.
J. Bot. 75: 864-886
Sato T, Sato S. 1984. Morphology of aecia of rust fungi. Trans. Br. mycol. Soc. 85: 223-238.
Sultan MA, Haq I, Khalid AN, Bajwa R. 2006. Some Uredinales from Northern areas of Pakistan.
Pak. J. Bot. 38 (3): 837-841.
Sultan MA, Haq I, Khalid AN, Mukhtar H. 2007. A contribution to Uredinales of Northern Areas
of Pakistan. Mycopath 4(2): 9-11.
Stewart RR. 1972. History and exploration of plants in Pakistan and adjoining areas. Flora of
Pakistan, pp. 7.
Wilson M, Henderson DM. 1966. British Rust Fungi, Cambridge Univ. Press, Cambridge, U.K.
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MY COTAXON
Volume 105, pp. 29-36 July-September 2008
A new species of /saria isolated from an infected locust
*ZONGQI LIANG’, XUEYOU HE’, YANFENG HAN’,
SHOUPING CAI’ & JIANGDONG LIANG!
zqliang1@yahoo.com.cn' swallow112886@yahoo.com.cn' ddl1977@yahoo.com.cn'
‘Institute of Fungus Resources, College of Life Sciences, Guizhou University
Guiyang, China 550025
hexueyou@public.fz.fj.cn? caishouping@gmail.com?
?Fujian Academy of Forestry, Fuzhou, China 350012)
Abstract — Isaria locusticola sp. nov. was isolated from a locust in Fuzhou, Fujian
Province, China. The new species is described and illustrated, and its relationships to
the related species are briefly discussed. This fungus is characterized by the variability
of the shape of conidia. Morphological characteristics and molecular analyses of ITS-
5.8S region sequences strongly support the establishment of I. locusticola in the genus
Isaria.
Key Words — hyphomycetes, Paecilomyces, entomopathogenic fungi, morphology,
rDNA analysis, Orthoptera
Introduction
Hodge et al. (2005) reviewed the nomenclatural status and history of the generic
name Isaria and concluded that Isaria Pers. is a valid name. Consequently, they
suggested that the name Isaria be used for those species previously assigned
to Paecilomyces Bainier section Isarioidea Samson. Luangsa-ard et al. (2005)
accepted this suggestion based on their molecular analysis and accepted 10
Paecilomyces species having Cordyceps teleomorphs as belonging in the genus
Isaria. These species were I. farinosa (Holmsk.) Fr., I. amoenerosea Henn.,
I. cateniannulata (Z.Q. Liang) Samson & Hywel-Jones, I. cateniobliqua (Z.Q.
Liang) Samson & Hywel-Jones, I. cicadae Migq., I. coleopterorum (Samson &
H.C. Evans) Samson & Hywel-Jones, I. fumosorosea Wize, I. ghanensis (Samson
& H.C. Evans) Samson & Hywel-Jones, I. javanica (Frieder. & Bally) Samson
& Hywel-Jones and I. tenuipes Peck. They also suggested that both P ramosus
Samson & H.C. Evans and P. xylariiformis (Lloyd) Samson belonged to Jsaria.
Corresponding author: Zonggqi Liang, zqliang1@yahoo.com.cn
30 ... Liang & al.
So far, more than 12 entomogenous species related to Paecilomyces have been
assigned to Isaria.
Species of Paecilomyces are considered the most important pathogens
parasitizing lepidoptoran insects (Evans & Samson 1982, Kubatova & Dvorak
2005). Species that parasitize orthopteran insects, however, have been rarely
reported. Evans was the first to isolate fungus P. reniformis Samson & H.C.Evans
from an infected grasshopper (Tettigoniidae, Orthoptera) (Samson 1974); the
same fungus has been also recently isolated from long horned grasshoppers
(Tettigoniidae, Orthoptera) in Sulawesi, Indonesia (Kalkar et al. 2006). Liang
et al. (1991) described Paecilomyces atrovirens Z. Q. Liang & A.Y. Liu from an
infected Tettigonia from Guizhou Province, China. During recent research on
Chinese entomogenous fungi, the authors discovered an undescribed Isaria
species from Fujian Province (eastern China), which is described below.
Materials and methods
Sample collection and strain isolation
Strain GZUIFR-Fj0916 was isolated from an infected adult of Chondracris sp.
(Acrididae, Orthoptera) collected in the Fuzhou National Forest Park, Fujian Province.
The infected insect collected from the field was first removed soil and other debris. Few
conidia were transferred to plates of Martins medium. Cultures were incubated at 26°C
until colonies were developed. Pure cultures having uniform conidiogenous structures
from the infected insect were transferred to Sabouraud’s slants and stored in the Institute
of Fungus Resources, Guizhou University.
Strain identification
The strain was transplanted to Czapek agar and potato dextrose agar (PDA),
incubated at 26°C for 14 days, and identified based on colony characters, conidiogenous
structures, temperature responses following Brown & Smith (1957) and Samson (1974)
and molecular analysis.
DNA extraction |
Taq enzyme and dNTP were supplied by Shanghai Sangon. ‘The strain GZUIFR-
Fj0916 used for the molecular identification was incubated on Czapek agar. DNA was
extracted from the fresh sporulating cultures following Tigano- Milani et al. (1995). The
extracted DNA was stored at -20°C.
PCR and ITS rDNA sequencing
The internal transcribed spacer (ITS) region including the 5.88 rDNA was amplified -
by polymerase chain reaction (PCR) using the primers ITS5 (5’- GGT GAG AGA
TTT CIG TGC -3’).and IT$4,(5-TGC TCC GCT TAT TGA TAT GG-3’). After a first
denaturation step at 94 °C for 5 min, the amplification reaction was performed for 35
cycles with denaturation at 94°C for 40 s, annealing at 49°C for 40 s, and extension at
Isaria locusticola sp. nov. (China) ... 31
Table 1 Fungi used in the study with their GenBank accession numbers
NAMES GENBANKNo NAME GENBANK No.
Byssochlamys fulva AY753341 I, javanica AY624186
B. nivea AY753339 I, javanica DQ403723
Cookeina colensoi AF394531 I. locusticola EU363501
C. venezuelae AF394041 Paecilomyces aerugineus AY753346
Cordyceps militaris AY725790 P. amoenerosea AY624169
C. takaomontana AY624198 P. carneus AY624170
Isaria japonica AF200370 P. marquandii AB099511
I. japonica AY624199 P. militaris EU363502
I. cateniannulata AF368802 P. niphetodes AY624192
I. cateniannulata AY624172 P. penicillatus AY624194
I. cateniobliqua AF368799 P. purpureus EF640809
I. farinosa AY624179 P. reniformis ARSEF429
I. fumosorosea AJ608982 P. reniformis ARSEF577
I. ghanensis AY624185 P. reniformis DQ069283
I. cateniobliqua AY624173 I. tenuipes AF368808
I. cicadae AB085887 P. variotii AY373941
_I. coleopterorum AY624176 P. viridis AY624197
72°C for 1 min; followed by a final extension step at 72°C for 10 min. PCR products were
purified using the Agarose Gel DNA Purification kit version 2.0 (Takara Company)
according to the manufacturer's protocol and were sequenced with the above primers
at Beijing Sunbiotech Co. Ltd. The sequence of ITS1-5.8S-ITS2 rDNA region of strain
GZUIFR-Fj0916 was submitted to GenBank (accession number: EU363501).
Sequence alignment and phylogenetic analysis
Table 1 lists strains used in the molecular study. Some ITS1-5.8S-ITS2 region
nucleotide sequences of representative Isaria and Paecilomyces species were obtained
from GenBank. The sequence of the new species was aligned with related fungi using
the Clustal X1.83 computer program for multiple sequence alignment and manually
corrected. The phylogenetic tree was constructed by neighbor-joining method (NJ) of
MEGA version 3.1 (Kumar et al. 2004). Confidence values for individual branches were
determined by bootstrap analysis (1000 replications).
Results and discussion
Taxonomy
Isaria locusticola Z.Q. Liang, X.Y. He & Y.F. Han, sp. nov. Fig. 1
MycoBank MB 511579; GENBANK EU363501
In agaro Czapekii, coloniae 40 mm diam, 14 diebus ad 26°C, ablae ad luteolum; reversum
luteolum. Hyphae septatae, hyalinae. Conidiophora erecta, septata, hyalina. Phialides
4.2-4.8 x 1.2-2 um, e basi inflata ellipsoida, claviformisa vel cylindricusa in collum
longum angustatae. Conidia continua, hyalina, levia, ellipsoidea, (2.4—)3-3.6(-4.2) x
(1.2-)1.5-1.8(-2) um, vel 4.8 x 3 um; fusiformia vel cylindricusa 3.5-4.5 x 1.2-1.5 um.
Teleomorphosis ignota.
32 ... Liang & al.
Fig. 1 The host insect and conidiogenous structures of I. locusticola.
1. A locust infected by I. locusticola; 2-5. The conidiogenous structures; 6. Conidia.
Holotypus GZUIFR-Fj016 isolatus, e Chondracris sp.(Acrididae, Orthoptera), Fuzhou regio,
Fujian provinca, China. IX, 2007, X.Y. HE et S. P CAL in Guizhou Univ, conservatur.
Cotony on Czapek agar, attaining a diameter of 40 mm within 14 days at 26°C,
flat, white to yellowish, villiform. Reverse yellowish. VEGETATIVE HYPHAE
hyaline, smooth-walled. CONIDIOPHORES septate, forming branches with
Isaria locusticola sp. nov. (China) ... 33
Table 2. A comparison of morphological characters between I. locusticola and related
species
Names Colony Conidia Hosts
Ff ihe, iP dab orcen cylindrical, pee sp.
4.2-7.2 x 2.4-3um (Acrididae, Orthoptera)
cylindrical to fusiform
Li : hit ;
javanica white 57 ae 417 in lepidopteran insects
I. locusticola white-yellowish ecae Sh etligs ae ee
2.4-4.2 x 1.2-2 um (Acrididae, Orthoptera)
fusiform to oblong-ellipsoid
3.9-4.9 X1.2-1.5, 1m
greyish-green, — reniform Grasshopper
P. reniformis ‘ eh ais
olive-green 4.5-5.5 x 2-3.5 um (Tettigontidae, Orthoptera)
phialides in whorls of 2 to 4. PHIALIDEs divergent, 4.2-4.8 x 1.2-2 um, consisting
of an ellipsoidal, cylindrical or clavate basal portion and a distinct thin neck.
Conrp1A hyaline, smooth-walled, variable, ellipsoid, (2.4-)3-3.6(—4.2) x (1.2-)
1.5-1.8(-2) um, a few up to 4.8 x 3 um; fusiform to oblong-ellipsoid, 3.5-4.5 x
1.2-1.5 um. Bi-celled rare.
TELEOMORPH: unknown.
DISTRIBUTION: Fujian Province, China.
MATERIAL EXAMINED: The holotype, GZUIFR-Fj0916, was isolated by X.Y He & S.P. Cai
from an infected cotton grasshopper Chondracris sp. (Acrididae, Orthoptera), Fuzhou,
Fujian Province, China, September, 2007, deposited in the Institute of Fungus Resources,
Guizhou University.
The two species of Paecilomyces known to infect locusts — P. reniformis and
P. atrovirens (Samson 1974, Liang et al. 1991, Kalkar et al. 2006) — distinctly
differ from I. locusticola (Table 2). P reniformis produces grayish-green or
olive-green colonies and reniform conidia, while Isaria locusticola produces
white colonies and ellipsoid to oblong-ellipsoid or fusiform conidia. Some
interesting bioassay tests also show that P reniformis isolated from grasshoppers
in Indonesia did not infect lepidopteran larvae at the concentrations tested and
the exposure method used, although the species did infect long- and short-
horned tettigoniid grasshoppers, suggesting that P reniformis may be specific
to this insect group (Kalkar et al. 2006).
Paecilomyces atrovirens differs from I. locusticola by its dark green colony,
long cylindrical conidia (4.2-7.2 um), phialides with a lecythiform basal
portion and imbricate conidial chain (Liang et al. 1991). I. locusticola and
I. javanica have similar conidiogenous structures , but the conidia of the former
are ellipsoid,(1.2-2 x 2.4-4.2 um), fusiform to oblong- ellipsoid (3.5-4.5 x
1.2-1.5 um), while they are cylindric (5-7.4 x 1.5-1.7 um) in the latter (Samson
34 ... Liang & al.
iG
~t
$6
£.98
Tsciria tenuipes AP 368808
* Cordyceps takaomontana AY624198
L. japonicadF 200370
id os BL japonicaAY624199
1. ghanensisAY624185
é 1. fumosoroseasJ6038982
I, coleopterorumAY624176
1, civcadaeABO8588?
I. cateniannulataAF 368802
88 § F cateniannulataAY624 172
ag jPaecilomyces militariskU363502
C. militar isAY7 25790
1 farinosaAY624179
P. reniformisDQ069283
P, reniformisARSEFS?7
P. reniformisARSEF429
eg [4 Javanicad Y624186
L javanicaDQ 403723
f locusticolakU 363501
L, amaeneraseaAY624169
1. cateniabliquaAY624173
I. cateniobliquadF 368799
P, niphetodes AY624 192
P. penicillatusAY624 194
P. carneusAY624170
P. purpureusEF 640809
P, viridisAY624197
P. marquandiABO995 11
P. aerugineusAY 733346
P. var totiiaY 373941
Byssochlamys filva A¥753341
Il-2
100
my
[l-2-1
36
-1
Byssochlamys ntvea AY733339
Cookeina colensoidF 394531
150 Cookeina venezuelaedF 394041
Fig. 2 Phylogenetic tree based on analysis of rDNA ITS1-5.8S-ITS2 sequences of I. locusticola
and some related species.
1974). A morphological comparison of I. locusticola and related species is
shown in Table 2.
Molecular identification
A BLAST search through GenBank was performed by using the ITS sequence of
I. locusticola as the query. Close matches showing maximal sequence identities of”
92-98% included I. javanica, I. fumosorosea, I. cateniannulata, I. cateniobliqua,
I. amoenerosea, and I. farinosa. The ITS sequences of these species and other
related species of Paecilomyces and other entomogenous fungi were retrieved
from GenBank for phylogenetic analysis.
Isaria locusticola sp. nov. (China) ... 35
Relationships of I. locusticola and related species in Isaria and Paecilomyces
were showed in the phylogenetic tree based on analysis of rDNA ITS1- 5.8S
- ITS2 sequences (Fig. 2). Cookeina colensoi and C. venezuelae were designated
as outgroups.
Luangsa-ard et al. (2004, 2005) demonstrated that Paecilomyces is
polyphyletic, with species falling in two subclasses, Sordariomycetidae
and Eurotimycetidae. Two clades (Clades I and II) were recognized in the
phylogenetic tree. Clade I consisted of the thermophilic Byssochlamys fulva,
B. nivea, P. aerugineus and P. variotii in the Eurotiales (Eurotimycetidae). Clade
II consisted of Paecilomyces spp. (Hypocreaceae, Hypocreales, Eurotimycetidae)
and Isaria spp. (Clavicipitaceae, Hypocreales, Eurotimycetidae,). I. locusticola
grouped with I. javanica as sister to I. cateniobliqua and I. amoenerosea. ‘This
association was well supported by bootstrap analysis (values = 96%).
Luangsa-ard et al. (2005) did not include P. reniformis in their phylogenetic
analyses. Our molecular analyses suggest that P reniformis does not group with
other Paecilomyces species but rather belongs to the entomogenous Isaria group,
which further suggests that P reniformis belongs to Isaria, not Paecilomyces.
Although both P reniformis and I. locusticola parasitize locusts, their separation
in the tree support them as independent species.
Both I. cateniobliqua and I. amoenerosea, which are characterized by red
colonies, are in the same subclade but apparently differ from the subclade of
I. locusticola and I. javanica. Although I. locusticola and I. javanica are closely
related by molecular analysis, they produce different conidia and live on
different insect hosts. Both the phenotypic description and molecular analyses
support recognition of strain GZUIFR-Fj0916 as a new member of Isaria.
Acknowledgements
This study was supported by the National Natural Science Foundation of China (No.
30499340). We are grateful to Prof. Y.C. Dai and Dr. R.T.V. Fox for their comments
on the manuscript. We also warmly thank Dr. $.R. Pennycook and Dr. L.L. Norvell for
editorial review and revisions.
References
Brown AHS, Smith G. 1957. The genus Paecilomyces Bainier and its perfect stage Byssochlamys
Westling. Trans Brit Mycol Soc. 40(1): 17- 89.
Evans HC, Samson RA. 1982. Cordyceps species and their anamorphs pathogenic on ants
(Formicidae) in tropical forest ecosystem I. The Cephalotes (Myrmicinae) complex. Trans Brit
Mycol Soc. 79: 431- 453.
Hodge KT, Gams W, Samson RA, Korf RP, Seifert KA. 2005. Lectotypification and status of Isaria
Pers.:Fr. Taxon 54 (2): 485-489.
Kalkar O, Carner GR, Scharf D, Boucias DG. 2006. Characterization of an Indonesian isolate of
Paecilomyces reniformis. Mycopathologia 161: 109-118.
36 ... Liang & al.
Kubatova A, Dvorak L. 2005. Entomopathogenic fungi associated with insect hibernating in
underground shelters. Czech Mycol. 57(3-4): 221-237.
Kumar S, Tamura K, Nei M. 2004. MEGA3: Integrated software for Molecular Evolutionary
Genetics Analysis and sequence alignment. Briefings in Bioinformatics 5: 150-163.
Liang ZQ, Liu AY, Feng DM. 1991. Some entomogenous fungi from Fanjing Mountain Preserve in
China. Acta Mycologica Sinica 12 (2): 110 -117.
Luangsa-ard JJ, Hywel-Jones NL, Manoch L, Samson RA. 2005. On the relationships of Paecilomyces
sect. Isarioidea species. Mycol. Res. 109(5): 581-589.
Luangsa-ard JJ, Hywel-Jones NL, Samson RA. 2004. The polyphyletic nature of Paecilomyces sensu
lato based on 18S-generated rDNA phylogeny. Mycologia 96 (4): 773-780.
Samson RA. 1974. Paecilomyces and some allied Hyphomycetes. Studies in Mycology 6: 1-119.
Tigano-Milani MS, Samson RA, Martins I, Sobral BWS. 1995. DNA markers for diferentiating
isolates of Paecilomyces lilacinus. Microbiology 141: 239-245.
MYCOTAXON
Volume 105, pp. 37-42 July-September 2008
Calathella columbiana (Basidiomycota):
new record of a cyphelloid fungus from Brazil
MARCELO A. SULZBACHER’, DENNIS E. DESJARDIN? & JAIR PUTZKE!
marcelo_sulzbacher@yahoo.com.br
'Universidade de Santa Cruz do Sul, Laboratorio de Botanica
Av. Independeéncia, 2293, Santa Cruz do Sul, RS, BRAZIL
ded@sfsu.edu
*San Francisco State University, Department of Biology
1600 Holloway Ave., San Francisco, CA, 94132, USA
Abstract — Calathella columbiana is described and illustrated based on material
collected in Santa Catarina State, southern Brazil. The species was originally described
from Colombia in 1983 and no more recorded from South America, the present record
is the second one for the species in this continent.
Key words — Agaricomycetidae, Cyphellaceae s.l., neotropical fungi
Introduction
Calathella D.A. Reid is a small genus of tubular or cupulate cyphelloid fungi
described by Reid (1964). Agerer (1983) studied six Calathella species and
considered the following characters to distinguish it from other related genera:
incrusted surface hairs with rounded tips, suburniform basidia, and oblong-
elliptical to cylindrical basidiospores. Currently, eight species are known in the
genus (Kirk et al. 2001).
From Brazil, there are few reports on cyphelloid mushrooms (Viégas
1945, Putzke 2002), possibly because of their much reduced size and difficult
taxonomy. The genus Calathella was only recently reported in Brazil by de
Meijer (2001, 2006), who mentioned unidentified specimens growing on
Cyathea tree-ferns.
In this paper, we report the occurrence of Calathella columbiana, the first
species reported from Brazil, collected in montane vegetation of Santa Catarina
State.
38 ... Sulzbacher, Desjardin & Putzke
Materials and methods
Specimens were collected and photographed, then macroscopically analyzed
following Bodensteiner et al. (2001). Microscopical observations were made
from material mounted in 3% KOH and Melzer’s reagent. Measurements of the
basidiospores do not include the apiculus; the shape was determined based on
Q (length-width ratio) values according to Largent et al. (1977). Color codes
used in the description of the species are those from Maerz & Paul (1930).
Specimens are deposited in the herbarium SMDB.
Taxonomy
Calathella columbiana Agerer, Mitt. Bot. Staatssamml. Miinchen 19: 192, 1983.
Fics. 1-4
BASIDIOMES 0.5—0.7 mm diam., scattered to clustered, with a short, stalk-like,
attenuated base when young, cupulate, unlobed but some becoming lobed in
age, with 2—5 broad finger-like lobes; margins slightly involute; external surface
woolly, pale-brown, becoming nearly white at the margin. HYMENIUM smooth,
glabrous, cream-colored. SuBicuLUM lacking. EXTERNAL SURFACE HAIRS 2—4
um diam., somewhat twisted, with obtusely rounded tips; tips hyaline, grading
into pale yellowish toward the base from intraparietal pigmentation; thickness
and color of adjacent trama hyphae similar to those of the bases of external
hairs; hairs with irregular incrustation patterns, with alternating densely
incrusted and nearly crystal-free areas, and tips sometimes loosely incrusted
or entirely glabrous; hairs not swelling in 5 % KOH, dextrinoid. TRAMA
neither agglutinated nor gelatinous; hyphae 2.5—4 um diam, with walls 0.5
um thick; outer tramal elements +/- parallel to basidiome surface, frequently
septate, yellowish from intraparietal pigmentation; SUBHYMENIAL HYPHAE
hyaline, short-celled, hardly differentiated from inner tramal hyphae. CLAMP
CONNECTIONS absent. Bastp1A 24—27 x 6—7 um, clavate, 4-spored; sterigmata
narrowly horn-like; lacking distinctive transition elements between basidia
and surface hairs at basidiome margin. BASIDIOLEs (17—) 19-27 (—32) x (3.5-)
5—7.5 um, clavate. CysTrp1a absent. BASIDIOSPORES 7.5—10 x (-3) 4—5 um [x =
8.4+0.7 x 4.1 + 0.6 um, Q= 1.7 - 2.7,Q_ =2.4 + 0.6, n = 20], asymmetrically
oblong-elliptical to cylindrical or slightly allantoid, apiculate, smooth, hyaline,
thin-walled, inamyloid, nondextrinoid.
HABITAT: scattered to gregarious on the abaxial surface of decayed grass leaves _
Poaceae, in cleared forest.
STUDIED MATERIAL: BRAZIL. Santa Catarina State, municipality of Riqueza, Linha Alta
Riqueza, (27° 02°01,4”S — 53°20°13,2”W), 340 m alt., 27.XII.2006, leg. A.A. Spielmann &
M.A. Sulzbacher-89 (SMDB 10.988)
Calathella columbiana in Brazil ... 39
Fig. 1. Calathella columbiana. Fresh basidiomes.
REMARKS: The studied Brazilian material is similar to C. digitiformis Bodenst.
et al. (Bodensteiner et al. 2001), another recently described species from Bali,
Indonesia. However, the latter species presents a set of features that differ from
C. columbiana,; i.e., digitiform basidiomes 0.7—1.5 mm diam. with numerous
finger-like lobes, narrower basidiospores (2.5—3.5 um diam. with mean width
3 um; Q_ = 2.8) and smaller basidia (14.5—21 x 4.5-6 um — Bodensteiner et al.
2001). Both species grow on herbaceous substrates.
The Brazilian collection exhibits intermediate characters between the species
cited above. For example, the size of the basidiospores and basidia are smaller
than those described for C. columbiana by Agerer (1983). In addition, a few
basidiomes are slightly lobed, but not as dissected as those of C. digitiformis.
Based on the neotropical distribution and macroscopic features, we consider
our specimen within the morphological variation of C. columbiana, which is
a poorly known species. Prior to this report C. columbiana was known only
from Colombia (type locality) and its distribution is now extended to southern
Brazil.
AO ... Sulzbacher, Desjardin & Putzke
Fig. 2-4. Calathella columbiana: 2) Basidiospores. 3) Basidia. 4) Basidiome section.
The taxonomic circumscription and phylogenetic position of the genus
Calathella is somewhat controversial, and recent molecular studies indicate
unresolved phylogenetic relationships suggesting that Calathella is not
monophyletic. Calathella columbiana was segregated from the other Calathella
species included in the analyses of Bodensteiner et al. 2004. In this work,
C. columbiana was sister of Mycenoporella lutea Overeem, a mycenoid species
with a poroid hymenophore and together they were sister of the Entolomataceae.
In comparison, C. mangrovei E.B.G. Jones & Agerer, and C. gayana (Lév.)
Agerer were distantly related to each other in the Nia clade and collectively
were sister of the Omphalotaceae. It should be noted, however, that the
relationships amongst the major clades presented in Bodensteiner et al. (2004)
were significantly different from those in the multi-locus analyses of Matheny
et al. (2006). Singer (1986) accepted Calathella as a synonym of the genus
Flagelloscypha because both taxa formed cyphelloid basidiomes covered with
dextrinoid, thick-walled hairs, and he maintained the distinction from the genus
Lachnella whose members have inamyloid hairs. In addition Singer (1986) was
Calathella columbiana in Brazil ... 41
reluctant to accept the numerous segregate genera recognized by Agerer (1983)
that were delimited by subtle micromorphological features. Recent molecular
data (Bodensteiner et al. 2004; Binder et al. 2005) support the acceptance of
Calathella, Flagelloscypha and Lachnella as distinct genera, but the relationships
amongst them are equivocal, and as already indicated, Calathella is polyphyletic.
It should be noted that the type species of Calathella, C. eruciformis (Batsch)
D.A. Reid, is not included in any published molecular studies, and accordingly
whether its phylogenetic affinities lie with C. columbiana or C. mangrovei or
C. gayana cannot be ascertained. Until the generic circumscription of Calathella
is clarified, we tentatively accept C. columbiana in Calathella.
Acknowledgments
The authors thank to Adriano A. Spielmann for the photograph of the collection. To
Vagner G. Cortez and Gilberto Coelho for providing the line drawings. To Dr. Timothy
J. Baroni (State University of New York - College at Cortland) and Dr. D. Jean Lodge
(Center for Forest Mycology Research, USDA Forest Service) for kindly reviewing the
manuscript.
Literature cited
Agerer R. 1983. Typusstudien an Cyphelloiden Pilzen IV. Lachnella Fr. s.1. Mitt. Bot. Staatssamml
Miinchen 19: 163-334.
Binder M, Hibbett DS, Larsson K-L, Larsson E, Langer E, Langer G. 2005. The phylogenetic
distribution of resupinate forms across the major clades of mushroom-forming fungi
(Homobasidiomycetes). Syst. Biodiv. 3: 113-157.
Bodensteiner P, Agerer R, Desjardin DE, Horak E. 2001. A new species of Calathella from Bali.
Mycologia 93: 1010-1013.
Bodensteiner P, Binder M, Moncalvo JM, Agerer R, Hibbett DS. 2004. Phylogenetic relationships of
cyphelloid homobasidiomycetes. Mol. Phylogen. Evol. 33: 501-515.
de Meijer AAR. 2001. Mycological work in the Brazilian state of Paranda. Nova Hedwigia 72:
105-159.
de Meijer AAR. 2006. Preliminary list of the macromycetes from the Brazilian state of Parana. Bol.
Mus. Bot. Munic., Curitiba 68: 1-55.
Kirk PM., Cannon PE, David JC, Stalpers JA. 2001. Ainsworth & Bisby’s Dictionary of the Fungi.
9" ed. Wallingford (UK): CABI.
Largent DL, Jonhson D, Watling R. 1977. How to Identify Mushrooms to Genus. III. Microscopic
features. Eureka (USA): Eureka Publishing.
Maerz AJ, Paul MR. 1930. A dictionary of color. New York (USA): McGraw-Hill.
Matheny PB, Curtis JM, Hofstetter V, Aime MC, Moncalvo J-M, Ge Z-W, Yang Z-L, Slot JC,
Ammirati JF, Baroni TJ, Bougher NL, Hughes KW, Lodge DJ, Kerrigan RW, Seidl MT, Aanen
DK, DeNitis M, Daniele GM, Desjardin DE, Kropp BR, Norvell LL, Parker A, Vellinga EC,
Vilgalys R, Hibbett DS. 2006. Major clades of Agaricales: a multilocus phylogenetic overview,
Mycologia 98: 982-995.
Putzke J. 2002. Agaricales (Fungos - Basidiomycota) pleurotdides no Rio Grande do Sul. I -
Anthracophyllum, Aphyllotus, Campanella, Chaetocalathus e Cheimonophyllum. Cad. Pesq., Sér.
Biol. 14: 45-66.
42 ... Sulzbacher, Desjardin & Putzke
Reid DA. 1964. Notes on some fungi of Michigan. I. “Cyphellaceae’. Persoonia 3: 97-154.
Singer R. 1986. The Agaricales in Modern Taxonomy. 4th ed. Koenigstein (Germany): Koeltz
Scientific Books..
Viégas AP. 1945. Alguns fungos do Brasil, VII-VIII: Cyphellaceae e Thelephoraceae. Bragantia >:
253-290.
MYCOTAXON
Volume 105, pp. 43-52 July-September 2008
A new species within the Gymnopus dryophilus complex
(Agaricomycetes, Basidiomycota) from Italy
ALFREDO VIZZINI', GIOVANNI CONSIGLIO’,
VLADIMIR ANTONIN? & MARCO ContTU*
‘alfredo. vizzini@unito.it
Dipartimento di Biologia Vegetale, Universita di Torino
Viale Mattioli 25, 10125 Torino, Italy
*giovanni.consiglio@fastwebnet. it
Via Ronzani, 61 - I 40033 Casalecchio di Reno, Bologna, Italy
-vantonin@mzm.cz
Moravian Museum, Department of Botany, Zelny trh 6
CZ-659 37 Brno, Czech Republic
* marcocontu@interfree.it
Via Traversa via Roma, 12 (I Gioielli 2) - 07026 Olbia, Sassari, Italy
Abstract — Gymnopus inexpectatus, collected from a coastal ecosystem in Italy on a
dead branch of Quercus ilex, is proposed as a new species. ITS and LSU rDNA sequences
place it in the G. dryophilus complex within sect. Levipedes subsect. Levipedes, where
its compact coralloid pileipellis structure and broadly ellipsoid spores (Q = 1.5) make
it unique. A Latin diagnosis, technical description, illustrations, and taxonomic
speculations on Gymnopus-Marasmiellus relationships are provided. Comparisons with
the European, North/South American and Indonesian species that represent the most
closely related species are also presented.
Key words — Collybia, gymnopoid/omphalotoid clade, Mediterranean Agaricales,
Omphalotaceae, taxonomy
Introduction
The genus Gymnopus (Pers.) Roussel (Agaricomycetes, Agaricales,
Omphalotaceae) includes saprotrophic (mainly litter decay) non-mycotrophic
agarics (Antonin & Noordeloos 1997, 2008) with white to cream-coloured
spore-prints, stipes that are not insititious but usually with basal mycelium and/
or rhizomorphs, rooting or rarely originating from a sclerotium, a cutis type
pileipellis composed of narrow, cylindrical hyphae or a complex structure of
inflated terminal elements (dryophila—structure), inamyloid, non-dextrinoid,
44 ... Vizzini & al.
acyanophilic basidiospores, and non-dextrinoid contextual hyphae. Gymnopus
is a character-poor genus, since many taxa diverge in only very subtle features,
such as basidiomata macromorphology, pileipellis structure, cystidia shape and
basidiospore size. This complicates species identification, often daunting from
a morphological perspective.
While studying mycobiota of the Natural Park of Rimigliano (S. Vincenzo,
Livorno, Tuscany, Italy), 120 hectares of sandy coastal ecosystem formed by
holm oak (Quercus ilex L.) and cork oak (Q. suber L.) woods and by a dense
mediterranean pine forest (Pinus pinea L., P. pinaster Aiton), specimens of an
unusual Gymnopus species have been found. After comparison with species
from Europe, (e.g. Vilgalys & Miller 1987a, Noordeloos 1995, Bon 1999,
Antonin & Noordeloos 1997, 2008), North and South America (e.g. Vilgalys &
Miller 1983, Halling 1983, 1996, 1997; Mata et al. 2006) and Asia (Indonesia,
Wilson et al. 2004), it is considered to represent a new species. It is described
and illustrated below.
Materials and methods
Macro- and micromorphology
Macroscopic characters were examined from fresh material. Microscopical
studies are based on dried material using a Leica DM 4500 B and an Olympus
BX50 light microscope with magnifications up to 1000 x. Observations were
made on mounts in the following reagents: Congo Red in 10% ammonia,
Cresyl-Blue in water, Melzer’s reagent and 3% KOH. Measurements are based
on observation of 30 basidiospores of two basidiomata (apiculus not included).
The following abbreviations have been used: E = the quotient of length and
width of the spores; Q = the mean value of E values in all collections studied; L =
number of entire lamellae; | = number of lamellulae between each pair of entire
lamellae. All examined material has been deposited and preserved in MCVE
(Museo Civico di Storia Naturale, Venice, Italy). Herbarium abbreviations
follow Holmgren & Holmgren (1998). Latin description of the new species has
been deposited in MycoBank.
DNA amplification and sequencing
The DNA from a dried basidiome was extracted using the DNA Plant
Minikit (Qiagen) and following manufactor’s instructions. The partial internal
transcribed rDNA spacer (ITS) and the D2 region of the large subunit (LSU)
rDNA were amplified through the polymerase chain reaction (PCR). The —
primer pairs ITS1F/ITS4 for ITS (White et al. 1990, Gardes & Bruns 1993) and
0061/NDL22 for LSU (van Tuinen et al. 1998, Kjoller & Rosendahl 2000) were
used both for PCR reactions and sequencing. The PCR products were purified
Gymnopus inexpectatus sp. nov. (Italy) ... 45
by the PCR purification kit and sequenced by DINAMYCODE S.r.l. Searches
for similar sequences allowing taxonomic identification were conducted using
the BLASTN algorithm available through the NCBI (http://www.ncbi.nlm.
nih.gov/blast/Blast.cgi). The two consensus sequences have been deposited in
GenBank and the relative accession numbers are listed with the collection.
Taxonomy
Gymnopus inexpectatus Consiglio, Vizzini, Antonin & Contu, sp. nov. FIGURES 1,2
MycoBank MB 511701
Pileus 25-30 mm latus, convexus vel plano-convexus, hygrophanus, ochraceo-brunneus,
maculis ferruginosis designatus, laevis, glaber. Lamellae (sub)distantes, L = 25-30, 1 = 2-6
(7), adnatae, cremeo-roseae. Stipes longus tam quam pilei diametron vel vix longior, 5-6
mm crassus, cylindraceus, insiticius, concolor cum pileo, longitudinaliter fibrillosus, basi
tomentosus. Caro cartilaginea. Odor nullus. Sapor mitis. Sporae 5-6 (7.0) x 3.2-4 (4.5) um,
E = 1.4-1.7, (late) ellipsoideae, ovoideae, tenuitunicatae, hyalinae, laeves, inamyloideae.
Basidia tetraspora, clavata. Cheilocystidia 27-45 x 5-8 um, lageniformia, subfusoidea
vel subcylindracea, saepe rostrata, saepe irregularia vel subcoralloidea. Lamellarum
trama ex cylindraceis vel fusoideis hyphis constituta. Pileipellis ex singulari specie
“dryophila-structurae” efformata, ex versiformibus, cylindraceis, clavatis, fusoideis vel
utriformibus, hyalinis vel pigmento parietali luteolo-brunneolo hyphis cum digitiformibus
propagationibus, constituta. Caulocystidia discretis cystidiis vel (plerumque) terminalibus
cellulis similia. Fibulae adsunt ubique.
Holotypus in loco Rimigliano dicto prope S. Vincenzo (Livorno, Italy), 2.XII.2006 lectus, leg.
G. Consiglio, E. Franceschini et G. Perdisa, in Herbario MCVE sub n° 16126 conservatur.
ETryMoLocy: from inexpectatus (Latin), meaning “unexpected, unforeseen’.
PILEUS 25-30 mm, convex to plano-convex, applanate with age, with slightly
involute margin, hygrophanous, not translucently striate when moist, brownish
ochre, rusty red spotted, pallescent to ochraceous cream, smooth, glabrous,
slightly tomentose at margin. LAMELLAE rather distant, subdistant, L = 25-30, |
= 2-6 (7), adnate, rosy cream, with entire to hardly uneven, concolorous edge.
STIPE of the same length, or slightly longer, than pileus diameter, 5-6 mm thick,
cylindrical, slightly broadened at base, insititious, stuffed, concolorous with
the pileus, longitudinally fibrillose, tomentose at base, without rhizomorphs.
ConrTEXT cartilaginous, concolorous with the external surfaces. SMELL none,
indistinct. TasTE mild.
BASIDIOSPORES 5-6 (7) x 3.2-4 (4.5) um, on average 5.9 x 3.8 um, E = 1.4-1.7,
Q = 1.5, (broadly) ellipsoid, ovoid, smooth, thin-walled, hyaline, inamyloid
(Fig. 2b). Basrp1a 20-25 x 6-7.5 um, 4-spored, clavate, often subcapitate.
BASIDIOLES 14-23 x 3-6 um, clavate, cylindrical. CHEILOCYSTIDIA 27-45 x 5—-
8 um, lageniform, subfusoid or subcylindrical, often rostrate, often irregular to
subcoralloid, thin-walled (Fig. 2a). PLEUROCYSTIDIA absent. HYMENOPHORAL
TRAMA consisting of cylindrical to fusoid, thin-walled, up to 13 um wide hyphae.
PILEIPELLIS a special form of “dryophila-structure’, composed of versiform,
46 ... Vizzini & al.
Fig. 1. Gymnopus inexpectatus (holotype). Basidiomata. Scale bar = 10 mm.
cylindrical, clavate, fusoid or utriform, + thin-walled, 30-62 x 8-15 um
elements, which are irregular, coralloid or with + digitate projections, hyaline
to pale yellowish-brownish in KOH (Fig. 2d); with parietal and fine epiparietal-
encrusting pigment. STIPITIPELLIS a cutis consisting of cylindrical, parallel,
thick-walled (up to 1 um), smooth, up to 7 um wide hyphae. CAULOCYSTIDIA
in the form of discrete cystidia or (mostly) terminal cells, 17-48 x 4.5-10 um,
adpressed to erect, cylindrical, clavate, regular, irregular to coralloid, + thin- to
thick-walled (Fig. 2c). CLAMP CONNECTIONS present in all tissues. CHEMICAL
REACTIONS, no part of the basidiome dextrinoid or amyloid and not staining
green in KOH.
HasitTat. On a little branch of Quercus ilex fallen on the soil.
COLLECTION EXAMINED — Italy, S. Vincenzo, Livorno, in the Quercus ilex forest of
Rimigliano: two basidiomes on a branch of Q. ilex, 2°! Dec. 2006, leg. G. Consiglio,
E. Franceschini et G. Perdisa (holotype MCVE 16126; GenBank accession numbers
EU622905 and EU622906).
Molecular results
The ITS and LSU rDNA D2 regions were amplified from the dried basidiome.
The size of the ITS and LSU rDNA fragments was 238 bp and 606 bp, respectively.
Searching for similar sequences in GenBank database pointed to taxa within
the Gymnopus dryophilus complex (Tab. 1).
Gymnopus inexpectatus sp. noy. (Italy) ... 47
Fig. 2. Gymnopus inexpectatus. Micromorphologic characters of the basidiomata. a. Cheilocystidia.
b. Basidiospores. c. Caulocystidia. d. Pileipellis elements (all from holotype). Scale bar = 20 um.
Discussion
This new species is well characterised by a unique combination of features.
Nevertheless its taxonomic relationships are problematic: our collection
combines characters such as a gymnopoid habitus (viz. pileus relatively fleshy,
not distinctly striate, lamellae usually not distant, stipe not filiform to hair-
like but cartilaginous), that suggest a close relationship with Gymnopus, with
characters (viz., coralloid pileipellis structure, lignicolous habit, insititious stipe)
reminiscent of several Marasmiellus s.]. species (Antonin et al. 1997, Antonin
& Noordeloos 1993, 1997, 2008). However, recent molecular analyses by Mata
et al. (2004, 2006) and Wilson & Desjardin (2005) note that the type species of
A8 ... Vizzini & al.
Gymnopus (G. fusipes), Micromphale Gray (M. foetidum) and Setulipes Antonin
(S. androsaceus) cluster within the same clade and that Marasmiellus Murrill
represents an artificial, polyphyletic genus, suggesting that some Marasmiellus
species should be included with Setulipes and Micromphale within Gymnopus.
Analyses of ITS and LSU rDNA sequences place our fungus in the Gymnopus
dryophilus complex, a group of species belonging to section Levipedes (Fr.)
Halling, subsection Levipedes (Antonin & Noordeloos 1997) (Tab. 1). This
subsection traditionally encompasses species that share the following anatomical
features: a pileipellis composed of relatively short, inflated, branched, loosely
arranged hyphae, usually not coralloid or diverticulate (forming the so-called
dryophila-structure, with elements which resemble a jig-saw puzzle, best seen
in paradermal scalps); well-differentiated cheilocystidia, and pileipellis hyphae
that do not turn green in alkali.
G. dryophilus could be conceived as a complex of morphologically similar
species. This complex was clearly distinguished from the rest of the section
Levipedes by both morphologically based phenetic and cladistic inferences
(Vilgalys 1986), and molecular analyses (Wilson et al. 2004, Mata et al.
2006). The presence of mating intersterile groups within this species complex
(“biological species”) from both North America (Vilgalys & Miller 1983)
and Europe (Vilgalys & Miller 1987b) was also supported by DNA/DNA
hybridization experiments (Vilgalys 1991). Concurrent morphological studies
also demonstrated small morphological differences among species recognized
within either continent (Vilgalys & Miller 1983, 1987a). These species and their
infraspecific taxa could be distinguished based on a combination of shape and
size of cheilocystidia together with rhizomorphs and gill colour.
Wilson et al. (2004) and Mata et al. (2006) are the first to attempt to
circumscribe the taxa within this complex using ITS sequences. Both Bayesian
and parsimony analyses of sequences within sect. Levipedes and the G.
dryophilus complex show little variation and differ from one another by only 5-
10 bp (Mata et al. 2006). Morphological plasticity and lack of resolution of ITS
sequence analyses complicate assignment of complex members to morphotaxa.
Mata et al. (2006) observe that the G. dryophilus complex seemingly has a Euro-
American distribution; this may be an artefact resulting from poor specimen
sampling.
Within this section and complex, G. inexpectatus is well characterized by (1)
a stipe with poorly developed basal mycelium, (2) growth on Quercus ilex wood
debris, (3) a trichoderm-like pileipellis comprising coralloid digitate elements,
irregular to subcoralloid cheilocystidia, and (4) broadly ellipsoid spores (Q = ©
1.5). These results show that sect. Levipedes (Fr.) Halling s. Antonin et al. (1997)
may also contain species with coralloid pileipellis cells as an extreme case of
dryophila-structure.
Gymnopus inexpectatus sp. nov. (Italy) ... 49
Table 1. Similarities of G. inexpectatus ITS and LSU rDNA sequences to those retrieved
from the GenBank database.!
ACCESSION # DESCRIPTIONS IDENTITY (%) E*VALUE
ITS
DQ480098 Gymnopus dryophilus voucher Duke 09 7e-114
DQ480097 Gymnopus dryophilus Duke 242 99 7e-114
DQ449960 Gymnopus ocior voucher TFB4284 2) 7e-114
DQ449959 Gymnopus ocior voucher TFB9015 99 7e-114
DQ449957 Gymnopus ocior voucher TFB3861 SPS) 7e-114
LOLA BE PE ah NE a ua al Ol nbd E@b waale
AF042595 Collybia dryophila isolate RV83/180 99 0.0
AY640619 Gymnopus dryophilus isolate AFTOL-ID 559 99 0.0
AF291305 Gymnopus dryophilus 28S large subunit rDNA, 99 0.0
partial sequence
AY639411 Gymnopus bicolor voucher AWW116-SFSU on 0.0
AF261328 Micromphale foetidum strain JEJ.VA.567 97 0.0
‘Search method: algorithm BLASTN 2.2.17 (Jun. 24, 2007) available at the NCBI site (http://www.ncbi.nlm.nih.
gov/BLAST/), performed on Febr. 8, 2008.
(*) The expected E value represents the estimated number of sequences randomly corresponding (the lower the
value, the better the correspondence with the sequence in the database).
None of the Gymnopus monographs from Europe (Antonin & Noordeloos 1997,
2008; Bon 1999), the Americas (Vilgalys & Miller 1983, Halling 1983, 1996,
1997; Mata et al. 2006), or Asia (Wilson et al. 2004) include a taxon close to the
new species proposed here. The most closely related described species seems to
be G. dryophilus var. lanipes (Malencon & Bertault) A. Ortega et al. (Malencon &
Bertault 1975, Lonati 1986, Bon 1999, Vila & Llimona 2002, Ortega et al. 2003,
Antonin & Noordeloos 2008), which exhibits similar habit and microscopical
features. However, it differs from G. inexpectatus by having a non-insititious
stipe, long stipe hairs, a more distinctly “jigsaw” (dryophila-like) pileipellis
structure, and different spore size (E = 1.6-2.3, Q =1.85; Ortega et al. 2003). G.
dryophilus var. lanipes typically grows in Mediterranean thermophilous forests
(Morocco, Italy and Spain) among vegetal debris (such as Quercus ilex, Pinus
spp., or, more rarely, Cistus spp). Ortega et al. (2003), who studied the holotype
(n° 4508) in Malencon’s Montpellier herbarium (MPU), notes that the holotype
material does not correspond to the taxon described in the protologue. It has
a pubescent stipe, but its spores are larger (9-11 x 4-5 um), the pileipellis is
a cutis of typical filamentous hyphae without a dryophila-structure, and the
basidiomata are brown-reddish: this collection may represent a taxon within
the G. terginus group. Because Malencon knew G. terginus very well, Ortega et
al. (2003) supposed that a designation of the material as holotype was surely an
unintentional error and selected a neotype from Iberian material that fitted the
original description in all aspects, keeping the original name.
50 ... Vizzini & al.
One recently described Spanish species, G. bisporus (J. Carbo & Pérez-De-Greg.)
J. Carbo & Pérez-De-Greg. [= G. catalonicus (Vila & Llimona) Vila & Llimona
2006, fide Antonin & Noordeloos 2008)], differs from G. inexpectatus in having
clavate cheilocystidia, mono- and bisporic basidia and spores (8)8.9-10.6(12)
x (3.5)4-4.9(5.5) um in size (Pérez-De-Gregorio & Carbé 2002, Antonin &
Noordeloos 2008).
Based only on macro- and micromorphological features, G. inexpectatus
could be mistaken for a Marasmiellus, as interpreted by Antonin & Noordeloos
(1993), of sect. Dealbati Singer, subsect. Quercini Singer (with coloured pileus;
Singer 1973, Desjardin 1997). In Europe (Antonin & Noordeloos 2008), the
species belonging to the M. carneopallidus/M. mesosporus/M. maas-geesterani/
M. maritimus group may look like G. inexpectatus. M. carneopallidus
(Pouzar) Singer, which has similar pileipellis structure and caulocystidia,
can be distinguished by larger spores (7.5-12.5 x 4.5-6.5 um) and broadly
clavate simple cheilocystidia. M. mesosporus Singer, with similar cheilo- and
caulocystidia, has larger spores (10.5-14.5 x 5-7.5 um) and larger basidia
(25-40 x 8-12 um) (Takehashi et al. 2007). M. maas-geesterani Robich &
E. Campo has larger spores [(9)10-—12(13.5) x 5-6.5(7.5) um], differently shaped
cheilo- and caulocystidia, and larger basidia (35-45 x 6.5-8.5 um) (Robich &
Campo 2000). M. maritimus Contu & Noordel (Noordeloos & Antonin 2008 =
Marasmiellus roseotinctus Contu & Noordel., nom. illeg., non Pegler) has larger
spores (10-14.5 x 4-6 um) and larger basidia (22-35 x 7-10.5 um) (Noordeloos
& Contu 2007).
In comparison with similar South-American species described by Singer
(1973), M. xerophyticus Singer has a smaller, 7-13 mm broad, brown pileus
with whitish margin, larger spores [(8)9-11 x (4.2)5-6(7) um], larger basidia
(19-36 x 6-7.5 um), and differently shaped cheilocystidia; M. aurantiorufescens
Singer has an ochraceous pileus that dries to rufous, longer and narrower
spores [(4.5)5.5-6.7 x 2.5-2.8 um], differently shaped cheilocystidia, and
growth on oak leaves; M. dryogeton Singer has a smaller (5-11 mm) ochraceous
brown to cinnamon brown pileus, larger spores (6.2-8.5 x 3-3.5 um), smaller
cheilocystidia (17-20 x 2-8 um), different caulocystidia, and grows on oak
leaves; M. enodis Singer has a brown to deep brown, radially sulcate pileus,
larger spores [6.5-9 x 2.5-4(4.5) um], differently shaped and smaller (22-35 x
7-9.5 um) cheilocystidia and caulocystidia in the form of Crinipellis-like hairs,
and grows on dicotyledonous wood; M. dendroegrus Singer has an ochraceous-
cinnamon, sulcate and striped pileus, larger spores (6-8.5 x 2.8-4.5 um), |
differently shaped cheilocystidia and grows on dead wood of dicotyledons; M.
quercinus Singer has a chestnut to deep cinnamon, small (6-10 mm) pileus,
differently shaped, single, ventricose to fusoid cheilocystidia, different stipe
covering, and grows on leaves (less frequently on small sticks and acorns) of
Gymnopus inexpectatus sp. nov. (Italy) ... 51
Quercus spp. The four last mentioned species are further distinguished by a
distinctly dark brown incrusted hypodermium.
Acknowledgments
We would like to thank Antonio Ortega (Departamento de Botanica, Facultad de
Ciencias, Universidad de Granada, Espafia) and Jordi Vila (Societat Catalana de
Micologia - Facultat de Biologia Departament de Biologia Vegetal, Unitat de Botanica,
Universitat de Barcelona, Espafia) for providing helpful suggestions. The research
of the third author was supported by the Ministry of Culture of the Czech Republic
(MK00009486201).
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MYCOTAXON
Volume 105, pp. 53-58 July-September 2008
Phellinus mori sp. nov.
(Basidiomycota, Hymenochaetales) from northern China
YU-CHENG Dar
*Corresponding author, yuchengd@yahoo.com
Institute of Microbiology, Beijing Forestry University
Beijing 100083, China
BaAo-Kat Cul
baokaicui@yahoo.com.cn
Institute of Microbiology, Beijing Forestry University
Beijing 100083, China
WAN-QIANG TAO
twq@bifb.gov.cn
Station of Forest Protection, Beijing Municipal Bureau of Landscape & Forestry
Beijing 100029, China
Abstract — A new resupinate member of the Hymenochaetaceae, Phellinus mori, is
described from northern China. It belongs to the Phellinus laevigatus complex, but
differs by strongly cracked basidiocarps when mature or dry, subparallel skeletals along
the tubes, presence of cystidioles, ovoid or subglobose basidiospores, and habitat on
Morus exclusively.
Key words — lignicolous fungi, poroid fungi, polypore, taxonomy
Introduction
During an investigation on lignicolous and poroid fungi in China, several
specimens were collected on Morus from Heilongjiang Province and Beijing
area in northern China. These collections are macroscopically similar to
Fomitiporia punctata (P. Karst.) Murrill by resupinate and cushion-shaped
basidiocarps, but they become strongly cracked when mature or dry. White
mycelial strands are common in old tubes. Microscopically there are many
hymenial setae and hyaline, fairly thick-walled, nondextrinoid and moderately
cyanophilous basidiospores. The collections belong to the Phellinus laevigatus
54 ... Dai, Cui & Tao
complex but differ sufficiently from all Bevioush described taxa to be described
as a new species.
Materials and methods
The studied specimens are deposited at the Herbarium of Beijing Forestry
University (BJFC) and the Herbarium of Institute of Applied Ecology, Chinese
Academy of Sciences (IFP). The microscopic procedure follows Dai (1999).
Sections were studied at magnification up to x1000 by using a Nikon Eclipse
E600 microscope and phase contrast illumination. Drawings were made with
the aid of a drawing tube. In presenting the variation in the size of the spores,
5% of measurements were excluded from each end of the range, and are given
in parentheses. In the text the following abbreviations are used: IKI = Melzer’s
reagent, IKI- = negative in Melzer'’s reagent, KOH = 5% potassium hydroxide,
CB = Cotton Blue, CB+ = cyanophilous, CB- = acyanophilous, L = mean spore
length (arithmetic average of all spores), W = mean spore width (arithmetic
average of all spores), Q = variation in the L/W ratios between the specimens
studied, n = number of spores measured from given number of specimens.
Special colour terms followed Petersen (1996) and Anonymous (1969).
Description
Phellinus mori Y.C. Dai & B.K. Cui, sp. nov. Figs.1-3
MycoBank MB 511704)
Carpophorum perenne, resupinatum. Facies pororum cinnamomeum vel brunneum, pori
rotundi, 7-8 per mm. Systema hypharum dimiticum, hyphae generatoriae septatae sine
fibulis, hyphae skeletales contextum 2-4 um in diam. Setae sdsunt in hymenio, 11.6-26
x 5-8.4 um. Sporae ovoideae vel subglobosae, pallidae, crassitunicatae, 4.5-5.2 x 3.8-4.6
um.
Type. — China. Heilongjiang Prov., Ning’an County, Jingbohu Nature Reserve, on living
tree of Morus, 8.1X.2007 Dai 8363 (holotype in IFP, isotypes in BJFC & H).
ETYMOLOGY — mori (Lat.): referring to the host tree genus.
Fruirnopy — Basidiocarps perennial, resupinate, firmly attached to the
substrate, not readily separable, without odour or taste when fresh, woody
hard and medium in weight when dry, up to 15 cm long and 6 cm wide, and
1 cm thick, usually receding and becoming cushion-shaped with age. Pore
surface cinnamon brownish when fresh, becoming dark brown when bruised,
clay buff and strongly cracked when mature or dry, with a slightly silky sheen;
sterile margin narrow to almost lacking, concolorous with pores; pores mostly ~
circular, some slightly sinuous, (6—)7—8(-—10) per mm (n=90/3); dissepiments
thin, entire. Context present at each layer, cinnamon brown to fawn brown,
hard corky, ca. 0.1 mm thick. Tubes concolorous with pores, woody hard; white
mycelial strands usually present in old tubes; tube layers distinct.
Phellinus morus sp. nov. (China) ... 55
Figs. 1-2. Phellinus mori. Fig. 1. Fresh basidiocarp. Fig. 2. Dried basidiocarp.
56 ... Dai, Cui & Tao
b-f:
10 pm
Fig. 3. Microscopic structures of Phellinus mori (drawn from the holotype). —a: Basidiospores.
__b: Basidia and basidioles. —c: Cystidioles. —d: Setae. —e: Hyphae from trama. —f: Hyphae from
context.
Phellinus morus sp. nov. (China) ... 57
HyPHAL STRUCTURE — Hyphal system dimitic; all septa without clamp
connections; tissue darkening but otherwise unchanged in KOH.
CONTEXT — Generative hyphae hyaline to pale yellowish, thin-walled, frequently
branched and simple septate, 2-3.2 um in diam; skeletal hyphae dominant,
rust brown, thick-walled with a narrow lumen to subsolid, unbranched, rarely
septate, flexuous, interwoven, 2.5-4 um in diam; hyphae of white mycelial tufts,
hyaline, thin-walled, strongly branched and flexuous, 1-1.8 um in diam.
TuBes — Generative hyphae hyaline, thin-walled, occasionally branched,
frequently septate, 1.5-2.5 um in diam; skeletal hyphae dominant, rust brown,
thick-walled with a narrow lumen to subsolid, more or less straight to flexuous,
subparallel along the tubes, 2.2-3.5 um in diam. Setae frequent, ventricose to
subulate, some furcate, dark brown, thick-walled, 11-24 x 5-8.5 um; fusoid
cystidioles occasionally present, 8.7-14.7 x 3.4—5.2 um; basidia barrel-shaped,
with four sterigmata and a simple septum at the base, 9-13.2 x 5.3-8 um;
basidioles in shape similar to basidia, but slightly smaller; secondary hyphae of
mycelial strands hyaline, thin-walled, frequently branched and septate, 1.5-2
ttm in diam. Rhomboid crystals frequently present in trama and hymenium.
SPORES — Basidiospores mostly ovoid, sometimes subglobose, hyaline, fairly
thick-walled, smooth, IKI-, moderately CB+, (4—)4.3-5.2(-5.4) x (3.5-)3.8-
4.6(-4.8) um, L=4.74 um, W=4.16 um, Q=1.11-1.31 (n=210/7).
ADDITIONAL SPECIMENS (PARATYPES) EXAMINED. — China. Beijing, Daxing County,
Gusangyuan, dead branch of living Morus, 4.[X.2007 Dai 8291, 8292, 8293, 8294, 8295,
8297, 8298, 8300, 8301, 8303 & 8305 (IFP & BJFC). Heilongjiang Prov., Ning’an County,
Jingbohu Nature Reserve, on living tree of Morus, 8.IX.2007 Dai 8309, 8310, 8339, 8342,
8344 & 8353 (IFP & BJFC).
Discussion
Phellinus mori belongs to the P. laevigatus complex, which includes P. betulinus
(Murrill) Parmasto, P laevigatus (P. Karst.) Bourdot & Galzin, P prunicola
(Murrill) Gilb., P rhamni (Bondartseva) H. Jahn, and P. spiculosus (W.A.
Campb. & R.W. Davidson) Niemela. Parmasto (2007) studied the complex and
provided statistical analyses of the basidiospores.
Phellinus mori resembles P. laevigatus s. str., but basidiospore shapes differ in
the two species: ovoid or subglobose in P. mori, while ellipsoid in P laevigatus
s. str. In addition, basidiospores of P. laevigatus s. str. (4.56-5.46 x 3.58-4.15
um, L=5.01 um, W= 3.87 um, Q=1.21-1.39, Parmasto 2007) are narrower than
those in P mori. Macroscopically, the basidiocarp of Phellinus mori is strongly
cracked when mature or dry, while it is not cracked in P. laevigatus s. str.
According to Parmasto (2007) Phellinus laevigatus occurs in Europe, West
Siberia and Middle Asia, and its hosts are exclusively trees of Betula. Phellinus
58 ... Dai, Cui & Tao
mori was found in Beijing and Heilongjiang in northern China, growing
exclusively on dead branches and living trees of Morus.
Phellinus mori is morphologically similar to P. betulinus, and the latter species
occurs in China (treated as P. cf. laevigatus, Dai 1999). But P. betulinus has
distinctly smaller basidiospores (3.25-4.55 x 2.51-3.49 um, Parmasto 2007),
and it has been found so far only on Betula.
Among the members of the Phellinus laevigatus complex, basidiospores of
P. prunicola, P. rhamniand P. spiculosus are 5.21-6.43 x 3.86—4.63 um, 5.46—-5.78
x 4.22-4.82 um and 4.99-6.1 x 3.83-5.23 um respectively (Parmasto 2007), and
P. mori differs from these species by smaller basidiospores. Basidiospores of
Phellinus spiculosus and P. mori may be slightly overlapped, but the former has
basidiocarps not cracked and lacks cystidioles. Ecologically Phellinus spiculosus
causes cankers on Quercus and Carya (Gilbertson and Ryvarden1987), but
P. mori does not cause cankers, and lives on Morus exclusively.
Acknowledgements
We are grateful to Drs. Mario Rajchenberg (Centro Forestal CIEFAP, Argentina) and
Zheng Wang (Yale University, USA) for reviewing the manuscript. The study is supported
by the Department of Beijing Science and Technology (Project No. D0705002040391)
and the Ministry of Science and Technology of China (Project No. 2005DFA30280).
Literature cited
Anonymous. 1969. Flora of British fungi. Colour identification chart. Her Majesty’s Stationery
Office, London.
Dai YC. 1999. Phellinus sensu lato (Aphyllophorales, Hymenochaetaceae) in East Asia. Acta Bot.
Fennica 166: 1-115.
Gilbertson RL, Ryvarden L. 1987. North American polypores 2. Fungiflora, Oslo. 452 pp.
Parmasto E. 2007. Phellinus laevigatus s.l. (Hymenochaetales): a ring species. Folia Cryptog.
Estonica 43: 39-49.
Petersen JH. 1996. Farvekort. The Danish Mycological Society’s colour-chart. Foreningen til
Svampekundskabens Fremme, Greve.
MYCOTAXON
Volume 105, pp. 59-64 July-September 2008
Perenniporia minor (Basidiomycota, Polyporales),
a new polypore from China
HONG-XIA XIONG
xiongxiongbao@126.com
Institute of Applied Ecology, Chinese Academy of Sciences
Shenyang 110016, China
Graduate University, Chinese Academy of Sciences
Beijing 100039, China
Yu-CHENG DAT
*Corresponding author, yuchengd@yahoo.com
Institute of Applied Ecology, Chinese Academy of Sciences
Shenyang 110016, China
BaAo-Kali CuI
baokaicui@yahoo.com.cn
Institute of Microbiology, Beijing Forestry University
Beijing 100083, China
Abstract — A new polypore, Perenniporia minor, collected from Changbaishan Nature
Reserve, northeastern China, is described and illustrated. It is a saprophyte growing
on fallen angiosperm branches. Macroscopically it differs from other species in the
genus by tiny and pileate basidiocarps. Unlike most species of the genus, P minor is
characterized microscopically by almost negative skeletal hyphae in Melzer’s reagent.
P. minor and P. truncatospora have more or less similar basidiospores, but the latter has
smaller pores, and its basidiospores are nondextrinoid.
Key words — lignicolous and poroid fungi, Polyporaceae, taxonomy
Introduction
Perenniporia Murrill is a cosmopolitan genus with around 90 species described
and accepted (Gilbertson & Ryvarden 1987, Ryvarden & Gilbertson 1994,
Hattori & Lee 1999, Decock & Figueroa 2000, Decock 2001a, b; Decock et al.
2000, 2001; Nufiez & Ryvarden 2001, Dai et al. 2002, Decock & Ryvarden1999a,
60 ... Xiong, Dai & Cui
b, 2000, 2003a, b). Twenty-three species in the genus have been recorded from
China (Dai et al. 2002, Cui et al. 2006, 2007), and most of them occur in the
tropics or subtropics. During an investigation on wood-inhabiting fungi
in northeastern China, a tiny polypore was found that after microscopical
examination turned out to represent an undescribed species of Perenniporia.
Materials and methods
The studied specimens are deposited at the Herbarium of Institute of Applied
Ecology, Chinese Academy of Sciences (IFP). Anatomy was studied, and
measurements and drawings were made from slide preparations stained
with Cotton Blue. Drawings were made with the aid of a drawing tube. The
microscopic routine used in the study followed Yuan et al. (2006). In presenting
the variation in the size of the spores, the 5% of the measurements at each end
of the range are shown in parentheses. The following abbreviations are used: IKI
= Melzer’s reagent, IKI- = negative in Melzer’s reagent, KOH = 5% potassium
hydroxide, CB = Cotton Blue, CB+ = cyanophilous, CB- = acyanophilous, L =
mean spore length (arithmetic average of all spores), W = mean spore width
(arithmetic average of all spores), Q = variation in the L/W ratios between
the specimens studied, n = number of spores measured from given number of
specimens. Special colour terms are from Petersen (1996).
Description
Perenniporia minor Y.C. Dai & H.X. Xiong, sp. nov. Fig.1
MycosBank MB 511703
Carpophorum annuum, solitarium, pileatum; facies pororum cremea, pori rotundi, 4-6
per mm. Systema hypharum dimiticum vel trimiticum, hyphae generatoriae fibulatae,
hyphae skeletales contextum 2.7-5 um in diam. Sporae pallidae, ellipsoideae vel truncatae,
crassitunicatae, dextrinoidae, CB+, 4.9-6.2 x 3.8-4.5 um.
Type. — China. Jilin Prov., Antu County, Changbaishan Nature Reserve, Huangsongpu,
alt. 1100 m, on fallen branch of Acer, 14.1X.2007 Dai 9198 (holotype in IFP, isotype in
BJFC). i
ETyMOLOoGy — minor (L.), referring to the tiny basidiocarp.
FRUITBODY — Basidiocarps annual, pileate, solitary, soft corky, without odour
or taste when fresh. Pileus semicircular to spathulate, projecting up to 0.8 cm,
1 cm wide, and 0.3 cm thick. Pileal surface cream to very pale buff when fresh,
smooth, indistinctly concentrically zonate, becoming cinnamon-buff to pale
brick, distinctly concentrically zonate or sulcate when dry. Pore surface cream
when fresh, becoming cinnamon-buff when dry; pores round, 4-6 per mm,
tube mouths fairly thick, entire. Context white when fresh, becoming cream
and corky when dry, up to 0.1 cm thick; with a very thin cuticle present at the
Perenniporia minor sp. nov. (China) ... 61
Fig. 1. Microscopic structures of Perenniporia minor (drawn from the holotype).
—a: Basidiospores. —b: Basidia and basidioles.
—c: Hyphae from trama. —d: Hyphae from context.
62 ... Xiong, Dai & Cui
upper surface, cinnamon-buff. Tubes concolorous with pore surface, darker in
contrast with context, hard corky, up to 0.2 cm long.
HYPHAL STRUCTURE — Hyphal system dimitic to trimitic; generative hyphae
with clamp connections; skeletal hyphae IKI-, or very weakly amyloid, CB+;
tissue unchanged in KOH.
CONTEXT — Generative hyphae infrequent, hyaline, thin-walled, occasionally
branched, 2.2-3.5 um in diam.; skeletal hyphae dominant, hyaline, thick-walled,
mostly with a narrow lumen, occasionally branched (trimitic-like), flexuous,
interwoven, 2.7—5 um in diam.
TUBES — Generative hyphae infrequent, hyaline, thin-walled, occasionally
branched, 2-3 um in diam.; skeletal hyphae dominant, hyaline, thick-walled
with a narrow lumen, occasionally branched, flexuous, interwoven, 2.5-4.3
um in diam. Hymenium usually collapsed, basidia usually infrequent, barrel-
shaped to pyriform, with four sterigmata and a basal clamp connection, 13-
16 x 4.5-6.5 um; basidioles in shape similar to basidia, but slightly smaller.
Rhomboid crystals frequently present in trama and hymenium.
SpoRES — Basidiospores ellipsoid, mostly truncate, hyaline, thick-walled,
smooth, usually bearing a small guttule, dextrinoid, CB+, (4.7—)4.9-6.2(-7) x
(3.6-)3.8-4.5(-4.7) um, L=5.36 um, W=4.12 um, Q=1.26-1.35 (n=96/3).
ADDITIONAL SPECIMENS (PARATYPES) EXAMINED. — China. Jilin Prov., Antu County,
Changbaishan Nature Reserve, Huangsongpu, alt. 1100 m, on fallen angiosperm branch,
29.X1.2007 Wei 3500 (IFP). Fusong Country, Lushuihe, Hongwei, on fallen angiosperm
branch, 28.X1.2007 Wei 3467 (IFP).
REMARKS — In the field Perenniporia minor resembles a juvenile Tyromyces
chioneus (Fr.) P. Karst., but the latter is readily differentiated microscopically by
its thin-walled, nondextrinoid basidiospores.
The new species differs from other taxa in the genus by its tiny and pileate
basidiocarp (less than 1 cm), and almost negative skeletal hyphae in Melzer’s
reagent. Perenniporia contraria (Berk. & M.A. Curtis) Ryvarden, P truncatospora
(Lloyd) Ryvarden, P ohiensis (Berk.) Ryvarden and P. ochroleuca (Berk.)
Ryvarden also have small basidiocarps when young, but these species are
perennial. In addition, P contraria has smaller basidiospores (3.4-4 x 2.5-3.1
um, Cui et al. 2006), and spores of P. ohiensis (12-14 x 7-8.8 um, Dai et al. 2002)
and P. ochroleuca (9-12 x 5.5-7.5 um, Dai et al. 2002) are distinctly bigger than
those of P minor. P. truncatospora resembles P. minor in basidiospore shape and
size (5.3-7.5 x 4-5 um vs. 4.9-6.2 x 3.8-4.5 tm), but the former has smaller
pores (6-8 per mm) and basidiospores that are negative in Melzer’s reagent.
Unlike most species of Perenniporia, skeletal hyphae of P minor are
nondextrinoid, but this character is not unique to the species. In fact, several
species of Perenniporia have non- to faintly dextrinoid skeletal hyphae, e.g.,
Perenniporia minor sp. nov. (China) ... 63
P. medulla-panis (Jacq.) Donk and P. ohiensis; however, basidiospores of these
species are dextrinoid and cyanophilous. We consider the spore Melzer’s and
Cotton Blue reactions as more important and consider them as key characters
for the genus.
Acknowledgements
We are grateful to Drs. Mario Rajchenberg (Centro Forestal CIEFAP, Argentina)
and Zheng Wang (Yale University, USA) for reviewing the manuscript. The study is
supported by the National Natural Science Foundation of China (Project No. 30771730)
and the Ministry of Science and Technology of China for research grant (Project No.
2005DFA30280).
Literature cited
Cui BK, Dai YC, Decock C. 2006. Two species of Perenniporia (Basidiomycota, Aphyllophorales)
new to China. Fungal Science 21: 23-28.
Cui BK, Dai YC, Decock C. 2007. A new species of Perenniporia (Basidiomycota, Aphyllophorales)
from China. Mycotaxon 99: 175-180.
Dai YC, Niemela T, Kinnunen J. 2002. The polypore genera Abundisporus and Perenniporia
(Basidiomycota) in China, with notes on Haploporus. Ann. Bot. Fennici 39: 169-182.
Decock C. 2001a. Studies in Perenniporia. Some Southeast Asian taxa revisited. Mycologia
93: 774-795.
Decock C. 2001b. Studies in Perenniporia (Basidiomycetes, Polyporales): African taxa 1. Perenniporia
dendrohyphidia and Perenniporia subdendrohyphidia. Syst. Geogr. PJ. 71: 45-51.
Decock C, Buchanan P, Ryvarden L. 2000: Revision of some Australasian taxa of Perenniporia
(Basidiomycetes, Aphyllophorales). Australian Syst. Bot. 13: 823-844.
Decock C, Figueroa H. 2000. Studies in Perenniporia, Navisporus ortizii,a synonym of Perenniporia
martius, and a note on Navisporus and Perenniporia in Cuba. Crypt. Mycol. 21: 153-162.
Decock C, Figueroa H, Ryvarden L. 2001. Studies in Perenniporia. Perenniporia contraria and its
presumed taxonomic synonym Fomes subannosus. Mycologia 93: 196-204.
Decock C, Ryvarden L. 1999a. Studies in neotropical polypores. Some coloured resupinate
Perenniporia species. Mycol. Res. 103: 1138-1144.
Decock C, Ryvarden L. 1999b. Studies in Perenniporia: Perenniporia detrita and its taxonomic
synonyms. Mycologia 91: 386-395.
Decock C, Ryvarden L. 2000. Studies in neotropical polypores 6. New resupinate Perenniporia
species with small pores and small basidiospores. Mycologia 92: 354-360.
Decock C, Ryvarden L. 2003a. Perenniporiella gen. nov. segregated from Perenniporia, including a
key to neotropical Perenniporia species with pileate basidiomes. Mycol. Res. 107: 93-103.
Decock C, Ryvarden L. 2003b. Studies in Perenniporia (Basidiomycetes, Perenniporiales)
Pseudopiptoporus chocolates comb. nov., a synonym of Perenniporia ahmadii, and a note on
Pseudopiptoporus. Nova Hedwigia 77: 199-211.
Gilbertson RL, Ryvarden L. 1987. North American polypores 2. Megasporoporia - Wrightoporia.
Fungiflora, Oslo. 434-885 pp.
Hattori T, Lee S. 1999. Two new species of Perenniporia described from a lowland rainforest of
Malaysia. Mycologia 91: 525-531.
64 ... Xiong, Dai & Cui
Nufiez M, Ryvarden L. 2001. East Asian polypores 2. Polyporaceae s. lato. Synopsis Fungorum
14: 165-522.
Petersen JH. 1996. Farvekort. The Danish Mycological Society's colour-chart. Foreningen til
Svampekundskabens Fremme, Greve.
Ryvarden L, Gilbertson RL. 1994. European polypores 2. Synopsis Fungorum 7: 394-743.
Yuan HS, Li J, Huang MY, Dai YC. 2006. Antrodiella stipitata sp. nov. from Heilongjiang Province,
northeastern China, and a critical checklist of polypores from the area. Cryptogamie Mycologie
272129,
Volume 105, pp. 65-78 July-September 2008
A taxonomic study of Heterodermia
(Lecanorales, Ascomycota) in South Korea
based on phenotypic and phylogenetic analysis
XIN Li WEI’”*, HENG Luo?*, YOUNG JIN KOH? & JAE SEOUN HuR”
*ishur1@sunchon.ac.kr
‘Key Laboratory of Systematic Mycology & Lichenology, Institute of Microbiology,
Chinese Academy of Sciences, Beijing 100101, China
*Korean Lichen Research Institute, Sunchon National University
Sunchon 540-742, Korea
Abstract — A study on taxonomy and phylogeny of Heterodermia in South Korea
is presented for the first time. Phenotypic analysis was based on morphological,
anatomical, and chemical characters, and phylogenetic analysis was based on nrDNA
ITS sequences. The results suggest that the genus Heterodermia is monophyletic. Some
characters (e.g. corticate vs. ecorticate lower surface, absence vs. presence of yellow
pigments or soredia) are shown to be taxonomically significant within the genus. Of
the eighteen species of Heterodermia known to occur in South Korea, eleven were
confirmed in this study. Brief species diagnoses accompany a key to Heterodermia in
South Korea, and comparisons with similar taxa are included.
Key words — lichen flora, macrolichen, Physciaceae, comprehensive analysis
Introduction
The genus Heterodermia belongs to the lichenized ascomycetous family
Physciaceae Zahlbr. Heterodermia is mainly characterized by linear lobes and
by spores of the Pachysporaria- or the Polyblastidia-type (Poelt 1965, Brodo
et al. 2001). All species of Heterodermia were included in Anaptychia Korb.
(Kurokawa 1962) until thick-walled spores and the presence of atranorin came
to be regarded as useful characters for separating these genera (Poelt 1965,
Swinscow & Krog 1976). Despite the many papers on Heterodermia (Awasthi
1973, Swinscow & Krog 1976, Wei 1991, Kurokawa 1998, Moberg & Nash 1999,
Grube & Arup 2001, Nordin & Mattsson 2001, Kurokawa 1962), almost no
studies on the genus were available for South Korea until the recent macrolichen
flora by Park (1990) that treated 12 Heterodermia species and provided brief
*Xin Li Wei and Heng Luo contributed equally to this work
66 ... Wei & al.
Table 1. Seventeen Heterodermia specimens used for DNA extraction
CoLL.# SPECIES NAME LOCALITY, GPS, ALT. (m) ACCESSION #
040340 H. boryi Mt. Chiri, N35°19’07.0” E127°40°42.7°, 1580 EU045422
040754 H. boryi Jeju.Do, N33 21 32,9" 8126 31 54.97 1920 EU045423
050093 H. diademata Mt. Deogyu, N35°51’24.1” E127°44’53.67,1580 EU045425
050273 H. diademata Mt. Sorak, N38°09718.1” E128°19°46.1, 880 EU045426
060639 H. dissecta Mt. Chiri, N35°19'01.0” E127°44’30.1", 1160 EU045424
040658 H. hypochraea Jeju Do, N33°24’26.6” E126°29°46.0", 645 EU045427
060232 H. hypoleuca Mt. .Chir,,N3517935.3° B127°32750.3,1390 EU045428
050017 ~—«-H. hypoleuca Mt. Naejang, N35°29°46.1” E126°53’56.7”, 600 EU045429
050317 H. japonica Mt. Sorak, N38°09711.2” E128°26'24.2”, 1265 EU045431
060291 H. japonica Mt. Chiri, N35°18'14” E127°34’10.4”, 1430 EU045432
060173 H. microphylla Mt. Chiri, N35°17'25.78” E127°31’59.7°, 1420 EU045433
060494 H. microphylla Mt.Deogyo, N35°51°10.0” E127°44°56.9, 1568 EU045434
050136 H. obscurata Wando, N34°21'10.3” E126°41°10.9", 535 EU045435
040623 H. obscurata Jeju Do, N33°28'25.5” E126°29°47.7°, 145 EU045436
050493 H. pseudospeciosa Mt. Haegsok, N34°41’21.1” E126°40°47.5” EU045438
040619 H. subascendens Jeju Do, 145, N33°28'25.5” E126°29°47.7” EU045441
040847 H, subascendens Jeju Do, 370, N33°27'15.4” E126°33’41.1” EU045442
descriptions of each species and a key to the taxa. However, some ambiguous
morphological characters left some taxonomic problems in need of additional
research. The most recently published checklist of Korean lichens (Hur et al.
2005), reports twenty Heterodermia species from the Korean peninsula. Thus,
in the present study, we have tried to clarify the taxonomy of Heterodermia
in South Korea based on specimens collected during recent nationwide field
surveys. In addition, we also evaluate whether Heterodermia is monophyletic
and discuss the importance of certain morphological and chemical characters
in differentiating species.
Materials and methods
MATERIALS — Two hundred and fifty-six specimens from South Korea have been
examined and are deposited in the herbarium of the Korean Lichen Research Institute
(KoLRI) in Sunchon, Korea. No type specimens were included in this study.
MorrHo.ocy — ‘The description of the external morphology is based on air-dried
material observed under a dissecting microscope. For anatomical descriptions, sections
were made with a razor blade under the stereomicroscope, mounted in GAW (glycerol:
alcohol: water = 1:1:1), and observed under a light microscope (Olympus BX 50).
Illustrations were taken with a digital camera (Nikon Coolpix 4500).
CHEMISTRY — Lichen substances were detected by the color reaction of a spot test (with
KOH) and by thin-layer chromatography (with B solvent system) (Culberson 1972,
White & James 1985).
Heterodermia in South Korea ... 67
Table 2. Sixty-nine phenotypic characters chosen for phenotypic analysis of
Heterodermia
No. CHARACTER No. CHARACTER
1 Thallus foliose 36 Yellow pigment present in the lower surface
2 Thallus subfiliform oe Rhizines marginal
3 Lobes linear 38 Rhizines at lower surface
4 Lobes subdichotomously branched 39 Rhizines marginal and at lower surface
>» Lobes irregularly branched 40 Rhizines black
6 Lobes discrete 4} Rhizines tan
7 Lobes crowded 42 Rhizines white
8 Lobe tips plane 43 Rhizines concolorous to thallus
9 Lobe tips ascending 44 Rhizines simple to squarrosely branched
10 Pape mnscitcinle 4s Paice dibs to subdichotomously
11 Lobe tips with pruina 46 Medulla white
12 The pruina of lobe tips round-patch 47 Pycnidia and (or) apothecia not seen
13 sti ie preps peace Get nlte 48 Pycnidia laminal, limited near tips
14 Lobe tips sometimes orange 49 Conidia bacilliform
15 Soredia present 50 Conidia small
16 Soredia marginal ba Conidia big
17 Soredia laminal Lys Apothecia lecanorine
18 Soredia marginal and laminal 53 Apothecia laminal
19 Soredia at lower surface 54 Apothecia terminal
20 Round-patch soralia 55 Apothecia marginal
21 Labriform soralia 56 Apothecia margin with lobules
22 Capitate soralia 57 The lobules of apothecia margin pruinose
23 Isidia present 58 Disks pruinose
24 Lobules present aS) Eight ascospores per asci
25 Lobules marginal 60 Ascospores brown
26 Lobules marginal and laminal 61 Ascospores 1-septate
27 Lobules tips suberect 62 Upper cortex prosoplectenchymatous
28 Lobules tips with pruina 63 Grayish surface layer present
og Lobules isidioid 64 Gonidia green
30 Upper surface maculate 65 Lower cortex prosoplectenchymatous
31 Upper surface papillate 66 No substances contained in the thallus
32 Lower surface corticate 67 Atranorin and zeorin present
33 Lower surface white 68 Norstictic acid present
34 Lower surface pale brown 69 Norstictic and salazinic acids present
35 Lower surface arachnoid
68 ... Wei & al.
PHENOTYPIC ANALYSIS — The phenotypic analysis is based on sixty-nine phenotypic
characters including morphological, anatomical, and chemical characters (Table 2).
Maximum parsimony analysis was performed using PAUP version 4.0b10 (Swofford,
2002). The reliability of the inferred tree was tested by 1000 bootstrap replications.
Anaptychia palmatula (Michx.) Vain. was used as outgroup for the analysis.
DNA EXTRACTION AND NRDNA AMPLIFICATION — Seventeen lichen thalli (Tablel) were
fractioned with cryo-tissue-crasher (SK200, Tokken, Japan). Total DNA was extracted
directly from whole thalli according to Ekman (1999) with DNeasy Plant Mini Kit
(QIAGEN, Germany), then purified by PCRquick-spin™ PCR Product Purification Kit
(iNtRON Biotechnology, INC.). ThenrDNAITS region (ITS1-5.8S-ITS2) was amplified by
PCR. The primers for amplification were: ITS1F (5'-CTTGGTCATTTACAGGAAGTAA-
3'; Gardes & Bruns, 1993) and ITS4A (5°-ATTTGAGCTCTTCCCGCTTCA-3;; White el
al. 1990). Conditions for PCR amplification and cycle sequencing have been described
previously (Arup 2002).
SEQUENCING AND PHYLOGENETIC ANALYSIS — PCR products were sequenced using
ABI 3700 automated DNA Sequencer in NICEM at Seoul National University. The
phylogenetic analysis was executed with software Mega3.1 (Kumar et al., 2004). The
Kimura 2-parameter model was selected, and gaps were retained initially while being
excluded in the pairwise distance estimation. The neighbor joining (NJ) (Saitou & Nei
1987) method was used to construct the phylogenetic tree, reliability of the inferred tree
was tested by 1000 bootstrap replications. Some Anaptychia species were available as
outgroups for the phylogenetic analysis.
Results and discussion
Phenotypic analysis
Phenotypic analysis (Fig.1) showed that all treated taxa of Heterodermia
form a monophyletic group, obviously separated from the outgroup Anaptychia
palmatula with 98% bootstrap support. Within Heterodermia, two sections
(I and II) based on the presence or absence of a lower cortex were indicated,
suggesting that the character-pair ‘lower surface corticate or ecorticate’ is
important to distinguish species.
Section I includes two species clusters (I-1 and I-2) according to the presence
of soredia and lobules, respectively. These characters are crucial in the taxonomy
among species with a lower cortex. In section IJ, no striking characters were
found to correspond to the topology structure except the spot test character of
the lower surface (K+, due to the presence of a yellow pigment). Three species
with this character — H. obscurata, H. hypochraea, H. subascendens — and
one species that sometimes expresses the character — H. japonica — clustered
in II-1 to some extent, suggesting that the spot test reaction has taxonomic
significance among species lacking a lower cortex. No other phenotypical
characters were shown to have taxonomic significance in Heterodermia.
Heterodermia in South Korea ... 69
Heterodermia japonica
53
Heterodermia obscurata
Heterodermia hypochraea Hi-1 Lower surface Kt;
51 yellow pigment present
Heterodermia subascendens
Il Lower surface ecorticate
Heterodermia boryi
Heterodermia hypoleuca
Heterodermia microphylla
98
98
Heterodermia diademata
|e Lobules present
Heterodermia dissecta
: : I Lower surface corticate
98 Heterodermia pseudospeciosa
: |u Soredia present
Heterodermia speciosa
Anaptychia palmatula
Fig. 1. Maximum parsimony tree based on morphological, anatomical and chemical characters
of 11 species of Heterodermia in Korea (Anaptychia palmatula = outgroup). Data matrix includes
12 taxa, 69 characters. All characters are of type ‘unord, all characters have equal weight, among
which 13 characters are constant, 14 variable characters are parsimony-uninformative, number
of parsimony-informative characters = 42. Tree length = 138, consistency index (CI) = 0.4058,
homoplasy index (HI) = 0.5942, CI excluding uninformative characters = 0.3387, HI excluding
uninformative characters = 0.6613, retention index (RI) = 0.4058, rescaled consistency index
(RC) = 0.1647. Heuristic search, bootstrap=1000. The numbers in each node represents bootstrap
support value, and the numbers lower than 50 were not shown.
Phylogenetic analysis
The NJ consensus tree (Fig.2) clusters all Heterodermia species into a well
supported (100% support) monophyletic clade that is clearly separated from
Anaptychia species. Two groups are included within the Heterodermia clade,
with group I species corticate and group II species ecorticate. This result agrees
with those from the phenotypic analysis and confirms that the presence or
absence of a cortical layer on the lower surface is an important diagnostic
character.
Within the group I, H. diademata, H. pseudospeciosa and H. dissecta
are clearly separated. On the one hand, H. diademata and H. dissecta differ
from H. pseudospeciosa by the absence or presence of soredia; on the other,
H. diademata differs from H. dissecta by absence or presence of norstictic and
salazinic acids, which are also present in H. pseudospeciosa. ‘This suggests that
absence or presence of soredia or norstictic and salazinic acids are important in
separating species in Group I.
70 ... Wei & al.
p A Heterodermia diademata EU045426
1001 A Heterodermia pseudospeciosa £U045438
A Heterodermia dissecta FU045424
, & Heterodermia hypoleuca EU045429
; A Heterodermia hypoleuca EU045428
100 | A Heterodermia microphylla EU045433
61> & Heterodermia microphylla EU045434
A Heterodermia obscurata EU045436
199! & Heterodermia obscurata EU045435 Group if lower cortex absent
100, A Heterodermia subascendens EU045441
1007! A& Hetorodermia subascendens EU045442
~
| Group I: lower cortex present
100, A Heterodermia diademata FU045425 |]
1
|
98 |
Monophyletic clade
~ & Heterodermia hypochraea £U045427 |
100,;-— A Heterodermia boryi FU045422
63] 1A Heterodermia boryi EU045423 |
A Heterodermia japonica EU045431 |
100! & Heterodenmia japonica EU045432 |
Anaptychia palmatala DQ394376
Anaptychia ciliaris AF389939
Anaptychia runcinata AF 224364
une Anaptychia bryorum AF2507&1
Anaptychia ulotricoides AF 540494
— Anaptychia elburzuana AY368 145
76
100
an |
0.05
Fig. 2. NJ consensus tree based on nrDNA ITS sequences. Nucleotide: Kimura 2-parameter,
pairwise deletion, bootstrap=1000. The numbers in each node represents bootstrap support value,
and the numbers lower than 50 were not shown. The sequences marked with Awere obtained in
this study, and others were downloaded from GenBank.
Group II shows species witha yellow pigment at the lower surface (H. hypochraea,
H. subascendens, H. obscurata) in one cluster separated from species lacking
this yellow pigment (H. boryi) or only sometimes possessing the pigment
(H. japonica), suggesting that presence or absence of yellow pigment may serve
as a helpful taxonomic character. Although there is only a short genetic distance
separating H. hypoleuca and H. microphylla, we think they should be treated
as two different species based on the phenotype showing two taxa are easily
distinguished phenotypically by rhizine color and the presence or absence of
soredia.
Based on our experimental results, we conclude that the genus Heterodermia
forms a well-supported monophyletic clade. We also identify four characters as
important for separating taxa within Heterodermia: (i) cortication of the lower
surface (Fig.3A, B), (ii) presence or absence of soredia (in some cases, Fig.3D),
(iii) presence or absence of a yellow pigment (Fig.3C), and (iv) the presence or
absence of norstictic and salazinic acids.
Taxonomy
Based on the comprehensive analysis given above, a key to the species is
presented, in which seven additional species not examined in our study
but reported before in South Korea are also included. All treated taxa are
subsequently listed and include diagnostic characters of each species. In case
Heterodermia in South Korea... 71
of large numbers of studied specimens only some representative specimens are
cited.
Key to the species of Heterodermia in South Korea
i ORS CATE ay <2) AS a a oh, Se Sr) A eee H. isidiophora
APASIC IA ADSCN Utes py hey ae ede etere pace es he ME an ce 2.
2, LOWenSUrlacercOrticatesss oa ia: jee aL NRC ACER aut Lae deca tt eee eee 3
Oe Lower surface noGeorticaté ainiiewe Ginn eRe eee eel tis ee ee eee a
mi ZINES Inapeitial Neate". eeeeaneree omer tt ear, sone cP: eee H. rubescens
Da IZINES NOM ar Cll alate cone ie ete. cyte eee es, oa eee cee nL ee er ae 4
A MLODESSOLCCIALC HORNS a) ie ates See Otis <p Caen ak tS | a ee D
A MICODES COOLCCIALE Lo atts Noligtt avis aucde.t eae meres Sams ao ek tol Ea tee aed ae ee 6
PR NOFStICLIGIACICH present <4iiut.. Mei nee rene nea ee H. pseudospeciosa
Da Norsticucacicabsent@. . aeiniee ys. 5 stip meet eanel ne cumin Proves tan, ete pe H. speciosa
6. Norstictic and salazinic acids absent; lobules marginal and laminal H. diademata
6. Norstictic and salazinic acids present; lobules only marginal............ H. dissecta
72 Lone andew nite, ciliapresentim: eft iraqake. Mat eee Genny sere eae ere H. comosa
Failte absentee. hei ae aa.” pies ban ad tee. lvo alse. 0c eae 8
SMBVEC CELL ARV CLO Wieeemites caCuis Naret een Nagle Wee rem © or leet kt ef ene H. firmula
SmIVICdULaWHItCte ee ee ee Cee eee ee 9
9. Lower surface K+ purple-red, yellow pigment present ......................-5. 10
DLowetsuttace Kesyellow pigmentabsent-a jacessiriacts ieee Seaiaases 15
TOMES esGrediaterts, 28102. el SMoiTi02 © Dit: .“olUGOl | LeOOoRL See noe lish
AVE ARCRAU Rel fota TEC Le tora Plas Lh ek AUREL) Pot Mn A robataetn Rio hed ah iy oo 9 oe 14
itt SiLowerrstirtace purplesblagk avcenters A0s ci. Ars Hee cite tat akira See Deen le
ivie WowelssUllace WhitesalCEN(Ch cage coc ts pineal concn Beene cae 13
12 URhizinestwiiteeeee weit ts ce er cae ose ot ROR Te ee aaa H. loriformis
PORE IZiN eS DAC Kee terme ie tame e. cos ne So skac car cee eee H. dendritica
13s Unknown yellowsupstalice present. a5. 920i) Ot, Ge eine 4 aa, H. pandurata
15, Unknownryellow substancembsent: sacs = ta. Oe tam See eee H. hypochraea
{AsSonediamtlower suriace non-labruormie orc ae ee H. subascendens
14 Leabriform. sora lian tie acc ste kat bekres cnet oby das ture cork ees eee H. obscurata
15: Thallus esorediatetn Wweahaeb es Face. igh ete seater eee H. hypoleuca
[5othallistsoretiate wt oe ee ee ec ee cs ete + COR SUE Ere ie See ave. 16
VOusGkecialaininalemixed WithslODUleSs ges. .e ee nn eer eee H. microphylla
LonSorediaimarginakorsséerminal 12.065 vive tlt yee eee nae ee 17
Wesercciaterminal thallus foliose. .utisn> sie ae cla ee eee eee eee H. japonica
ivPsoredia marpinalsthallus'subfiliform yee Oe a ee get H. boryi
72. ... Wei & al.
Heterodermia Trevis., Atti Soc. Ital. Sci. Nat. (Milano) 11: 613. 1868
Lectotype: H. speciosa (Wulfen) Trevis. (selected by Poelt 1965).
1. Heterodermia boryi (Fée) K.S. Singh & S.R. Singh, Geophytology 6 (1): 33, 1976
REMARKS: H. boryi is easily characterized by its strap-shaped to subfiliform lobes,
circinate tips, and the round or patch-shaped soralia that are lateral and laminal.
SPECIMENS EXAMINED: KOREA. Mt. Jiri, N35°19°09.1”, E127°40726.9”, alt. 1620 (1564)
m, on rock, 23 April 2004, Hur 040329; Mt. Jiri, N35°19°07.0” E127°40°42.7°, alt.
1580(1531) m, on rock, 23 April 2004, Hur 040340; Jejudo, N33°21°32.9" E126°31°54.9",
alt. 1920 m, on rock, 27 August 2004, Hur 040754.
2. Heterodermia diademata (Taylor) D.D. Awasthi, Geophytology 3: 113, 1973
REMARKS: H. diademata is characterized by a sublinear thallus, the abundance of
marginal and laminal lobules, and a corticate lower surface (Fig. 3B). The species is
similar to H. dissecta in morphology, but differs in chemistry, salazinic and norstictic
acids absent in H. diademata, while present in H. dissecta.
SELECTED (OF 30) SPECIMENS EXAMINED: KOREA. Mt. Naejang, alt. 680 m, on rock, 29
June 2003, Hur 030429; Baekdam temple, N38°11°16.4” E128°21’42.7°, alt. 450 m, on
rock, 11 October 2004, Hur 041509; Mt. Jiri, N35°20731.5” E127°41°08.9”, alt. 715 m, on
bark, 4 September 2004, Hur 040928; Mt. Deogyu, N35°51'24.1” E127°44’53.6", alt. 1580
m, on bark, 3 April 2005, Hur 050093; Mt. Sorak, N38°09'18.1” E128°19°46.1°, alt. 880 m,
on bark, 16 June 2005, Hur 050273; Wando arboretum, N34°21710.3” E126°41°10.9”alt.
535 m, on rock, 13 April 2005, Hur 050133.
3. Heterodermia dissecta (Kurok.) D.D. Awasthi, Geophytology 3: 113, 1973
REMARKS: H. dissecta is characterized by sublinear lobes, the abundance of marginal
(occasionally laminal) lobules, and a corticate lower surface. The species is similar to
H. diademata in morphology, but differs in chemistry, with salazinic and norstictic acids
present in H. dissecta and absent in H. diademata.
SPECIMENS EXAMINED: KOREA. Mt. Baega, N34° 10°14’, E127°8°45’, alt. 500 m, on rock,
8 October 2005, Hur 050572. Mt. Chiri, N35°19°1”, E127°44’30.1", alt. 1160 m, on rock,
15 September 2006, Hur 060639.
4. Heterodermia hypochraea (Vain.) Swinscow & Krog, Lichenologist 8 (2): 119,
1976
REMARKS: H. hypochraea has a papillate upper surface, erect lobe tips, and an ecorticate
lower surface with a yellow pigment (Fig. 3C). The species is similar to H. hypoleuca in
morphology, but differs in its white rhizines, erect lobe tips, and yellow lower surface.
SPECIMEN EXAMINED: KOREA. Jejudo, N33°24'26.6” E126°29°46.0’, alt. 645 m, on bark,
26 August 2004, Hur 040658.
5. Heterodermia hypoleuca (Mihl.) Trevis., Atti Soc. ital. Sci. nat. (Milano) 11: 615,
1868
Remarks: H. hypoleuca has a maculate upper surface, an ecorticate lower surface
(Fig. 3A), center that is reticulately veined, and hyaline to tan rhizines. This species
Heterodermia in South Korea... 73
Fig. 3. A. Ecorticated lower surface of Heterodermia hypoleuca, Hur 030174; B. Corticate
lower surface of H. diademata, Hur 030429; C. Yellow pigment (arrow) at the lower surface of
H. hypochraea, Hur 040658; D. Labriform soralia of H. pseudospeciosa, Hur 030204.
is similar to H. hypochraea, but differs by the tan colored rhizines, the plane lobe tips,
and the absence of a yellow pigment in the lower surface. H. hypoleuca is also similar to
H. japonica in morphology, but differs in that H. hypoleuca has hyaline to tan rhizines
and lacks soredia.
SELECTED (OF 107) SPECIMENS EXAMINED: KOREA. Mt. Baekwoon, alt. 1050 m, on
rock, 5 May 2003, Hur 030174; Mt. Hanla, alt. 1300, on moss, 22 June 2003, Hur 030333;
Mt. Sobaek, N36°56’50.5” E128°29°51.2”, alt. 1110 m, 2 October 2003, Hur 030752;
Mt. Gaya, N35°48’28.5” E128°07°00.6", alt. 965(916) m, on rock, 15 April 2004, Hur
040196; Mt. Odea, N37°43’42.8” E128°35736.6’, alt. 650(662) m, on Abies, 7 May 2004,
Hur 040400; Jejudo, N33°25’21.9” E126°33’35.7°, alt. 505(614)m, on bark, 26 August
2004, Hur 040676; Mt. Taebaek, N37°06700.3” E128°57702.5”, alt. 1220 m, on bark, 12
September 2004, Hur 041046; Mt. Naejang, N35°29°46.1” E126°53°56.7’, alt. 600 m, on
bark, 8 January 2005, Hur 050017; Mt. Deogyu, N35 51-292” E127 4503.4, alt 1530
m, on bark, 2 April 2005, Hur 050079; Mt.Chiri, N35°17°35.3” E127°32’50.3”, alt. 1390
m, on Quercus, 17 June 2006, Hur 20060232.
6. Heterodermia japonica (M. Sat6) Swinscow & Krog, Lichenologist 8 (2): 122,
1976
REMARKS: H. japonica is characterized by ascending lobe tips, labriform to subcapitate
soralia, an ecorticate lower surface, and the presence (‘H. propagulifera’) or absence
7A ... Wei & al.
(H. japonica s.str.) of a yellow pigment. This species is similar to H. hypoleuca, but differs
in having black rhizines and labriform to subcapitate soralia at the lobe tips.
SELECTED (OF 46) SPECIMENS EXAMINED: KOREA. Mt. Baekwoon, alt. 705 m, on bark, 1
June 2003, Hur 030282; Mt. Taebaek, N37°06’21.7” E128°57'12.1yalt. 1025 m, on rock,
2 November 2003, Hur 030841;Jejudo,33°24’26.6” E126°29°46.0", alt. 645 m, on bark,
26 August 2004, Hur 040654; Mt. Sorak, N38°06'40.4” E128°24’26.8’, alt. 1335 m, on
Abies, 10 October 2004, Hur 041460; Mt. Deogyu, N35°47°19.4” E127°42°30.3°, alt. 1269
m, on rock, Hur 050207; Mt.Chiri, N35°18'14” E127°34710.4”, on 1430 m, on rock, 17
June 2006, Hur 060291.
7. Heterodermia microphylla (Kurok.) Skorepa, Lichenologist 8 (2): 132, 1976
Remarks: H. microphylla has labriform soralia and the abundance of erect lobules (mixed
with granular soredia) on the upper surface. This species is similar to H. hypoleuca, but
differs in having labriform soralia, marginal and laminal lobules, and granular soredia
and lobules mix o the upper surface.
SELECTED (OF 36) SPECIMENS EXAMINED: KOREA. Mt. Naejang, alt. 695 m, on rock, 29
June 2003, Hur 030427; Mt. Taebaek, N35°29°46.9” E126°53’40.7°, alt. 1025 m, on rock, 2
November 2003, Hur 030837; Jejudo, N33°21732.3” E126°31°23.5”, alt. 1920 m, on bark,
27 August 2004, Hur 040762; Mt. Jiri, N35°17°25.78” E127°31°59.7°, alt. 1420 m, on
Quercus, 17 June 2006, Hur 060173; Mt.Deogyo; N35°51°10.0” E127°44°56.9", alt. 1568
m, on bark, 10 August 2006, Hur 060494; Mt.Baewoon, N35°36’31.3” E127°38°25.67,
alt. 1140 m, on Quercus, 17 August 2006, Hur 060605; Mt.Songni, N36°32’39.7”
E127°51°49.3”, alt. 700 m, on rock, Hur 060048.
8. Heterodermia obscurata (Nyl.) Trevis., Nuovo Giorn. Bot. Ital. 1: 114, 1869
REMARKS: H. obscurata has labriform soralia, marginal lobules, yellow pigmented lower
surface, and by black rhizines. Similar to H. subascendens, but differs in the occurrence
of a yellow pigment (which covers almost all the lower surface), the black rhizines, and
the striking labriform soralia at the lobe tips.
SPECIMENS EXAMINED: KOREA. Sorokdo, alt. 15 m, on bark, 23 March 2003, Hur
030034; Mt. Gaya, alt. 400 m, on bark, 22 April 2003, Hur 030153; Jejudo, N33°28'25.5”
E126°29°47.7°, alt. 145 m, on bark, 26 August 2004, Hur 040623; Jejudo, N33°25’21.9”
E126°33'35.7°, alt. 505(614) m, on bark, 26 August 2004, Hur 040680-1; Jejudo,
N33°27'15.4” E126°33’41.1’, alt. 370 m, on bark, 29 August 2004, Hur 040863, 040867;
Mt. Wonlak, on bark, 18 September 2004, Hur041171; Wando arboretum, 34°21710.3”
E126°41°10.9", alt. 535 m, on rock, 13 April 2005, Hur 050136; Mt. Hegsog, N34°41’21.4”
E126°40°51.4", alt. 203 m, on rock, 23 September 2005, Hur 050462.
9. Heterodermia pseudospeciosa (Kurok.) W.L. Culb., Bryologist 69: 484, 1967
REMARKS: H. pseudospeciosa is characterized by labriform soralia, lateral branches with
subcapitate soralia, and a corticate lower surface. The species is similar to H. speciosa .
in morphology, but differs in chemistry. Norstictic and salazinic acids are present in
H. pseudospeciosa, while absent in H. speciosa.
SPECIMENS EXAMINED: KOREA. Mt. Jiri, alt. 515 m, on rock, 4 May 2003, Hur 030204;
Mt. Baekwoon, alt. 700 m, on rock, 1 June 2003, Hur 030281; Mt. Kem, N34°45’32.3”
Heterodermia in South Korea... 75
E127°59°31.9", alt. 340(310) m, on rock, 15 Febuary 2004, Hur 040033; Baegdam temple,
N38°1116.4” E128°21°42.7”, alt. 450 m, on rock, 11 October 2004, Hur 041508; Mt.
Hegsog, N34°41'21.4” E126°40’51.4", alt. 203 m, on rock, 23 September 2005, Hur
050451; Mt. Hegsog, N34°41’21.1” E126°40°47.5”, alt. 230 m, on rock, 23 September
2005, Hur 050493; Mt. Juwang, N36°23’58.9” E129°09’53.1”, alt. 310 m, on rock, 16
October 2005, Hur 050618; Mt.Songni, N36°32’41.6” E127°51719”, on Quercus, 10
August 2006, Hur 060025.
10. Heterodermia speciosa (Wulfen) Trevis., Atti Soc. Ital. Sci. Nat. Milano 11: 614,
1868
REMARKS: H. speciosa has capitate soralia on the tips of lobes and lobules, and a
lower cortex. The species is similar to H. pseudospeciosa in morphology, but differs
in chemistry. Norstictic and salazinic acids are absent in H. speciosa, while present in
H. pseudospeciosa.
SPECIMENS EXAMINED: KOREA. Jejudo, alt. 145 m, on bark, 26 August 2004, Hur
040615; Jejudo, N33°28'25.5” E126°29°47.7’, alt. 145 m, on bark, 26 August 2004, Hur
040620; Jejudo, N33°27°15.4” E126°33’41.1’, alt. 370 m, on bark, 29 August 2004, Hur
040849; Mt. Juwang, N36°24’09.6” E129°10’27.0”, alt. 380 m, on rock, 15 October 2005,
Hur 050602.
11. Heterodermia subascendens (Asahina) Trass, Folia cryptog. Estonica 29: 20,
1997
REMARKS: H. subascendens is characterized by its maculate upper surface, marginal
and grayish white rhizines, an ecorticate yellow lower surface and granular soredia near
lobe tips. The species resembles H. hypoleuca in morphology, but differs in the color of
rhizines, the presence of a yellow pigment, and the presence of granular soredia on the
lower surface. It is also similar to H. hypochraea, differing in the color of rhizines and the
presence of granular soredia on the lower surface.
SPECIMENS EXAMINED: KOREA. Sorokdo, alt. 15 m, on bark, 27 December 2003, Hur
030895; Jejudo, N33°28725.5” E126°29°47.7”, alt. 145 m, on bark, 26 August 2004, Hur
040619; Jejudo, N33°27°15.4” E126°33’41.1°, alt. 370 m, on bark, 29 August 2004, Hur
040847.
Earlier reports on further species, not found in the present study
An additional seven species previously recorded from the Korean peninsula
but not found in our recent surveys are included in the key according to
descriptions provided in the literature.
1) Heterodermia comosa (Eschw.) Follman & Redon, Willdenowia, 6: 446, 1972
Its striking diagnostic character is the presence of long and white cilia on the lobe edges.
In Korea, the species has been recorded only once at a single locality on Jeju Island
by Kashiwadani et al. (2002). It is widely distributed in tropical regions of Asia and
America (Kurokawa 1962).
76 ... Wei & al.
2) Heterodermia dendritica (Pers.) Poelt, Nova Hedwigia 9: 31, 1965
The species is characterized by absence of soredia, often pruinose margins, lower surface
ecorticate, white, purple-black only at center, partly yellow pigmented. The taxon
is similar to H. propagulifera, but differs in the absence of soredia. Kurokawa (1962)
included H. propagulifera as a variety in H. dendritica (as Anaptychia). In Korea, this
species has been once recorded from Mt. Jiri by Park (1990).
3) Heterodermia firmula (Nyl.) Trevis., Atti Soc. Ital. Sci. Nat. Milano 11: 615, 1868
The diagnostic character of this species is the yellow medulla. It is close to the endemic
Indian species H. albidiflava (Kurok.) D.D. Awasthi, which shows a PD+ deep-yellow
reaction (unknown substance). In Korea, H. firmula was recorded only once from Mt.
Sobaek by Park (1990). However, we found no species with a yellow medulla on this
mountain, although we have many specimens collected there.
4) Heterodermia isidiophora (Nyl.) D.D. Awasthi, Geophytology 3: 114, 1973
Heterodermia isidiophora is a peculiar species in having corticate lower surface
and isidia in upper surface. In Korea, the species has been recorded by Park (1990).
However, during our study, all the similar specimens with isidia are Anaptychia species,
containing no chemical substances, so, whether H. isidiophora really exists in South
Korea still needs further study.
5) Heterodermia loriformis (Kurok.) Swinscow & Krog, Lichenologist 8: 129, 1976
Heterodermia loriformis is a peculiar species in having a white ecorticate lower surface
with zones of a yellow pigmentation and white rhizines. Similar to H. dendritica, but
differs in its white rhizines. In Korea, the species has been recorded only once by Park
(1990) as H. cf. loriformis based on specimens from Mt. Jiri. However, Moon (1999)
identified the specimens as H. pandurata. Further studies are still needed to clarify the
identity of the taxon.
6) Heterodermia pandurata (Kurok.) J.C. Wei, An enumeration of lichens in China,
p. 112, 1991
This species was reported by Moon (1999) on Mts. Jiri and Kaya of Korea. It is very
similar to H. hypochraea, but differs in presence of unknown yellow substance
(Kurokawa 1962). We didn't find this species during our study.
7) Heterodermia rubescens (Rasanen) D.D. Awasthi, Geophytology 3: 114, 1973
The species is characterized by a corticate lower surface and by mainly marginal
rhizines. It had been known only from India until Moon (1999) reported it from Korea. -
The voucher specimens are deposited in TNS (Japan), and we found no corresponding
specimens in our study.
Heterodermia in South Korea ... 77
Acknowledgments
This work was supported by the Korea Science and Engineering Foundation (KOSEF)
grant through Korean National Research Resources Center program (R21-2007-000-
10033-0). We are indebted to Prof. Bruce McCune and Dr. Walter Obermayer for giving
valuable comments on the manuscript. The first author also gives her thankfulness to
Prof. Jian-Bin Chen, Ms. Mei-Rong Ren, Kwang-Mi Lim and Hae-Sook Jeon for their
kind help and cooperation during this study.
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Volume 105, pp. 79-87 July-September 2008
Scutellospora striata sp.nov., a newly described
glomeromycotan fungus from La Gran Sabana, Venezuela
GISELA CUENCA! & RICARDO HERRERA-PERAZA2‘*
*gcuenca@Ivic.ve
‘Instituto Venezolano de Investigaciones Cientificas (IVIC), Centro de Ecologia
Apartado 20632, Caracas 1020-A, Venezuela
*Instituto de Ecologia y Sistematica, CITMA
AP 8029, Capdevila, Boyeros, La Habana, Cuba
Abstract — Examination of soil samples from a sclerophyllous shrubland near the
town of Iborib6 in La Gran Sabana, Venezuela, revealed an undescribed species
of Scutellospora whose spores are ornamented with finger-print-like processes, are
ochraceous yellow with a pinkish tint, and have a complex wall structure. The new
species, named Scutellospora striata, is the third Scutellospora species described from
La Gran Sabana, Venezuela.
Key words — arbuscular mycorrhizal fungus, AMF, Gigasporaceae, taxonomy, tropical
species
Introduction
The La Gran Sabana region is located in southeastern Venezuela, South
America. It is a highly undulate plain occupying ca. 30.000 km? of a slope from
1450 m to 750 m above sea level directed from the north to the south. This
mosaic of igneous and metamorphic rock is one of the earliest shields of the
earth crust. It formed the western section of the ancient supercontinent of
Gondwana (Schubert & Huber 1989). It was formed in Precambrian age and
due to its antiquity and advanced phase of weathering, soils originated from that
substrate (Roraima Formation) are mainly sandy with a very small clay fraction
and generally very low nutrient content. La Gran Sabana is also the land of the
tepuis, which are mountains of vertical walls and flat hilltops (tablelands), and
an outstanding region due to the high degree of endemism of its flora (Rull
1991). The plant cover of La Gran Sabana is an intricate mosaic, composed of
+Dr. Herrera-Peraza passed away in December of 2006.
80 ... Cuenca & Herrera—Perazat
numerous types of vegetation. With the exception of the continuous forests at
the foot of the eastern tepuis, forests occur in patches encircled by extensive
treeless savannas, as well as by shrublands (Dezzeo 1994).
Shrublands are primary plant communities in which the shrub stratum
constitutes the principal functional compartment. They consist of a well-
developed shrubby stratum 1-3 m high, occasionally with emergent shrubs up
to 5 or 7 m tall, with predominantly sclerophyllous leaves (Huber 1994), which
almost always grow on rocky sandstone outcroppings or on white quartzitic
sands. According to Huber & Febres (2000) they contain a significant number
of endemic plants. :
During the inventory of the arbuscular mycorrhizal fungi (AMF) associated
with the sclerophyllous shrublands of La Gran Sabana, spores of an undescribed
species of the genus Scutellospora with a distinctive ornamentation were found.
The fungus is described here as S. striata sp. nov.
Material and methods
Intermittent soil samples were taken during two years below vegetation ina sclerophyllous
shrubland situated at Iborib6 (5°36.796’N, 61°29.351’W) in La Gran Sabana. This
shrubland grows on sandstone outcrops (Huber 1994). It is highly diverse in plant
species, but among the most common are Clusia pusilla, Gongylolepis benthamiana,
Euphronia guianensis, Humiria balsamifera, Calliandra sp., Bonnetia sessilis and some
small rosettes belonging to the Cyperaceae family such as Rhynchospora barbata and
Bulbostylis conifera. Open pot trap cultures started from these soils were maintained
in a glasshouse for three months and then dried for one week. The content of the pot
was stored at room temperature (~20°C) for almost one year to break latency of spores
(Morton et al. 1993). Then, spores were isolated and used to start pure cultures that all
failed. Therefore, the description of the species presented below was prepared based
on spores isolated only from trap cultures and field soils. The cultures were initiated
with Vigna luteola as the host plant, but it died and then was replaced by wild plants
germinating from the seed bank. Various very interesting species of Scutellospora spp.
were isolated from this place (Herrera-Peraza et al. 2001, Walker et al. 1998). However,
no pure culture of any of them could be obtained.
Spores were isolated from the trap pots or from the field soils by wet sieving, decanting
and sucrose centrifugation (Sieverding 1991). The isolated spores were suspended in
water and illuminated with light from a quartz-iodine fibre-optic source. Their color
was determined by comparison with a color chart for British fungi (Anon 1969). The
specimens were mounted in polyvinyl alcohol lacto-glycerol (PVLG) or in PVLG
mixed with Melzer’s reagent (1:1, v/v). Wall description and terminology are based
on those suggested by Walker (1983) and Walker & Vestberg (1998). Type material has
been deposited in the Venezuelan National Herbarium (VEN) and the Cuban National
Herbarium, IES-CITMA (HAC).
To study more carefully the external appearance of the spores, a number of them
were prepared for Scanning Electron Microscopy (SEM). Prior to the preparation of
the material for SEM, spores were put under a dissecting microscope to remove or to
Scutellospora striata sp.nov. (Venezuela) ... 81
break the outer unit component with the aid of fine tweezers. Then, intact or broken
spores were rinsed in a phosphate buffer solution and immediately fixed in 1% osmium
tetroxide at 4°C for 1 h. Fixed samples were dehydrated for 5 min in each dilution of an
acetone series (20, 40,70,80 and 100 %), dried at critical point with liquid CO,, placed on
aluminum metal holders, coated with 200 A gold-palladium and observed for SEM.
Taxonomic description
Scutellospora striata Cuenca & R.A. Herrera sp.nov. Figures 1-3
MycoBAnk MB 511702
Spore singulatim enatae in solo, ochraceae roseo suffusae, superficie nitida, vulgo globosae
vel subglobosae, 130-184 x 116-152 um, supra suspensorem bulbosum ochraceum locatae.
Sporae parietum structura 6-tunicata. Tunicae prima et secunda respective unitaria
et laminata, caeterae omnes membranaceae. Tunica prima 1.5-2 crassa, laevis. Tunica
secunda 5-10 um crassa ochracea, in juventute tenuiter laminata et ornata muris 1.0-1.5
um latis, invicem 0.5-0.8 um separatis, striatim dispositis, vestigia digitalia referentibus.
Tunica tertia, 0,7 um crassa, rigida, firmiter tunicae secundae adpressa. Tunica quarta,
quinta et sexta, flexibles, membranaceae. Tunica quarta difficilis visu, sed Melzeris reactivo
dilute rosea evadens. Tunica quinta et sexta firmissime adpressae, solum sub microscopio
Normanski obviae, vel sexta Melzeris reactivo purpurea evadens. Scutellum germinativum
varie et dense plicatum. Cellulae auxiliares nobis non observatae.
Ho.LotyPe: Slide no. Cuenca 479-4, 13 Feb. 1998, Venezuelan Institute for Scientific
Research.
EryMo_oey Latin striata “with striae’, referring to the spore ornamentation composed
of elongated, generally parallel elements separated by grooves.
Spores borne singly in soil, ochraceous yellow of a pinkish tint in color
(Ochraceous [8G] to Saffron [49] according to the color chart), having a
shinning surface when observed under a dissecting microscope and illuminated
with tungsten lamp, generally globose or subglobose (Fig. 1a), 130-184 x 116-
152 um (mean 163 x 152 um, n= 13) with a terminal attached bulbous base,
produced from a septate subtending hypha. Bulbous spore base ochraceous
yellow in color and 22-29 um wide, with a recurved septum at its base.
SPORE WALL STRUCTURE composed of 6 components or layers generally
organized in 3 groups. With the exception of components | and 2, being unit
and laminated, respectively, all the remaining components are membranous.
Component 1 is contiguous with the sporophore outer wall component, 1.5-
2.0 um thick, and have a smooth surface (Figs. 2a, b and 3). Component 2 is
laminated, 5.0-10.0 um thick, and ochraceous yellow (Fig. 3). In young spores
the component 2 is finely laminated and ornamented with muri 1.0-1.5 um
wide and spaced 0.5-0.8 um apart (Fig. 1b). When seen in a plane view, the
muri resemble a finger-print. When spores develop 1 to 3 thick laminae (1.0-
2.0 um thick) originate from component 2 outwards (Figs. 3a-c). Each of these
laminae have the same ornamentation as described above. When three laminae
82 ... Cuenca & Herrera—Perazat
are formed, next fine inseparable laminae are synthesized inward. (Fig. 3c).
As mentioned before, each thick lamina is ornamented on the upper surface
with muri showing a striate pattern. The muri are commonly squared, angular
or wave-like in a cross section. The striations rarely follow the same direction
on the surface of each of the thick laminae and, thereby, in a plane view the
spore ornamentation seems to be composed of tiny squares or rhombs (Fig.
1c). At each two-muri interception wall material is deposited to form a small
columella. Therefore, in a cross section the component 2 is composed of up to
three lines of rounded cavities resulting from inter-muri and columella inter-
spaces (Fig. 2c).
Component 3, measuring approx. 0.7 tm, is a rigid membrane tightly
adhered to the lower surface of component 2 or separates from it in vigorously
crushed spores (Fig. 1h). Components 4 to 6 are flexible membranes usually
tightly adherent to each other.
Component 4 is generally very difficult to discern (Fig. 1f), but its presence
becomes evident in Melzer’s reagent because it reacts pale pink (Fig. 1g).
Components 5 and 6 are firmly cemented in between and can be discerned only
under a microscope equipped with Nomarski interference contrast or in spores
crushed in Melzer’s reagent because component 6 becomes readily purple in it.
None of these innermost membranes are beaded or amorphous. Germination
shield 64 x 72 um in size with complex infolding (Figs. le and 3b).
BULBOUS SPOROPHORE lacks ornamentation. The wall of the sporophore is
composed of two layers that are contiguous with components | and 2 of the
spore wall (Figure 1d). This wall is 1.5-2.0 um thick, reaching up to 3.0-5.0
um near the spore base. The pore at the attaching point is approximately 1.0
uum diam. Hyphae attached to the sporophore are regularly septated, measuring
5.0-9.0 um diam., and have walls 1.0-2.0 um thick. Auxiliary cells unknown.
DISTRIBUTION AND HABITAT. Known only from La Gran Sabana, Venezuela.
Spores of this species have been collected only from the sclerophyllous shrubland
at Iboribo and in a treeless savanna dominated by Axonopus canescens, also in
the way to the Iborib6 shrubland. Soils are highly acidic, sandy with a very low
exchangeable phosphorus and medium level of total N.
MYCORRHIZAL ASSOCIATIONS unknown. Attempts to form mycorrhizae in pure
culture have failed, though the species sporulated in a multispecies pot culture
with Scutellospora spinosissima, and other unknown species of Acaulospora,
Glomus, and Scutellospora.
COLLECTIONS EXAMINED: Type. Isolated from trap culture at the Venezuelan Institute
for Scientific Research, Caracas, Venezuela. Holotype slide no. Cuenca 479-4, 13 Feb.
1998. Isotypes: Cuenca 411-17, March 1997, Cuenca 386-1, 19 Dec. 1996.
Scutellospora striata sp.nov. (Venezuela) ... 83
Figure 1. Scutellospora striata. a) Intact spore mounted in PVLG. b) Ornamentation of spore
wall component 2 seen in a plane view. c) Ornamentation of spore wall component 2 of older
spore. It resembles rhombs due to the overlapping laminae, each ornamented with striae.
d) Non-ornamented bulbous suspensor. e) Germination shield. f) Crushed spore with inner group
of membranous components. Most internal components are labeled. g) Heavily crushed spore
mounted in PVLG+Melzer’s reagent with stained components 4 and 6. h) Cross view of spore with
components 1-6 arranged in two groups.
84 ... Cuenca & Herrera—Perazat
Figure 2. SEM images of S. striata. a) Intact spore with the outermost unit component obscuring
the ornamented spore wall component 2. b) The striae on the upper surface of component 2 visible
through the broken component 1. c) Cross view of component 2 with lines and round cavities.
Discussion
Scutellospora striata can be readily distinguished from other species in the
genus by its peculiar ornamentation. Under a dissecting microscope, the
ornamentation is not always evident, because the unit outer component is
smooth and usually shines under incident light, though it detaches from the
spore easily after low pressure. The presence of the very robust component 1
was evidenced by SEM (Fig. 2a). It has a smooth surface, which is often free of
organic debris. However, in some spores lacking the outer component 1, fine
soil debris firmly adhere to the spore surface, which show the ornamentation
when detached. The ornamentation of component 2 could be visualized under
SEM only after manipulating the spore to break the outer unit component (Fig.
2b).
Component 2 is quite complex. Because each layer of the laminated
component 2 is ornamented with striae, in a plane view the striae of overlapping
layers are visible as tiny squares or rhombs (Fig. 1c). When spores are very
young and only one layer (subcomponent) of component 2 is differentiated or
when the outermost subcomponent is in focus, the ornamentation resembles a
finger-print (Figs. 1b and 3a).
Assessing the number of wall components is very difficult. The number of
inner components may be determined when first intact spores are mounted
in PVLG for 1-2 hours and crushed by applying pressure to the cover slip to
Figure 3 (on next page). Differentiation of the laminate spore wall component 2 of S. striata.
a) Only one lamina of component 2 of young spore. Outer unit component 1 is transparent and
easy to see. b) Component | and two laminae of component 2 of older spore. Germination shield is
also visible. c) Component | and four laminae of component 2 of mature spore.
Scutellospora striata sp.nov. (Venezuela) ... 85
86 ... Cuenca & Herrera—Perazat
push the internal membranes out (Fig. 1f). Component 4 may also be revealed
in spores crushed in PVLG+Melzer’s reagent, because it stains light pink (Fig.
1g).
The sporulation of S. striata in both trap cultures and the field was not
abundant. Unfortunately, attempts to grow this fungus in one-species cultures
failed and, hence, the properties of its mycorrhizae remain unknown, similar
to those of many other unculturable members of the Glomeromycota (Fitter
2005).
Of the described species of the genus Scutellospora, eleven produce
ornamented spores. However, while the ornamentation of S. striata spores
consists of striae resembling a finger-print when seen in a plane view, that
of spores of S. coralloidea, S. dipapillosa, S. gregaria, S. heterogama, S. persica
and S. verrucosa is composed of warts, S. crenulata of dome-like subpolygonal
papillae separated by pits, S. minuta of spines with round apices, S. nigra of pits,
S. nodosa of knobs, and S. spinosissima of blunt spines.
In addition to S. crenulata and S. spinosissima, S. striata is the third newly
described species from La Gran Sabana, Venezuela.
Acknowledgments
Fundacite Guayana is gratefully acknowledged for the financial support provided.
Professor Bruno Manara from the Venezuelan Botanical Institute prepared the Latin
diagnosis and Milagros Lovera, Eduardo Furrazola and Rigel Fernandez gave invaluable
comments to the manuscript. We thank Dr. Janusz Blaszkowski and Dr. Marta Cabello
for reviewing the manuscript.
Literature cited
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Chart. Edinburgh Her Majesty’s Stationery Office.
Dezzeo N. 1994. Ecologia de la Altiplanicie de la Gran Sabana (Guayana Venezolana) I. Scientia
Guaianae 4: 1-205.
Fitter AH. 2005. Darkness visible:reflections on underground ecology. Journal of Ecology 93:
231-243.
Herrera-Peraza RA, Cuenca G, Walker C. 2001. Scutellospora crenulata a new species of Glomales
from La Gran Sabana Venezuela. Canadian Journal of Botany 79: 674-678.
Huber O. 1994. Los arbustales. Pages 95-106. in N. Dezzeo editor. Ecologia de la Altiplanicie de La
Gran Sabana (Guayana Venezolana) I. Scientia Guaianae N° 4.
Huber O, Febres G. 2000. Guia ecoldgica de La Gran Sabana. The Nature Conservancy Caracas.
192 pp. p
Morton JM. 1995. Taxonomic and phylogenetic divergence among five Scutellospora species based
on comparative developmental sequences. Mycologia 87: 127-137.
Morton JB, Bentivenga SP, Wheeler WW. 1993. Germ plasm in the international collection of
arbuscular or vesicular-arbuscular mycorrhizal fungi (INVAM) and procedures for culture develop-
ment documentation and storage. Mycotaxon 48: 491-528.
Scutellospora striata sp.nov. (Venezuela) ... 87
Rull V. 1991. Contribucién a la paleoecologia de Pantepui y la Gran Sabana (Guayana Venezolana)
clima biogeografia y ecologia. Scientia Guaianae N° 2:1-132.
Schubert C, Huber O. 1989. The Gran Sabana. Panorama of a Region. Lagoven booklets. Refolit
Venezuela.
Sieverding E. 1991. Vesicular-arbuscular mycorrhiza management in tropical agrosystems. GTZ
Eschborn Germany.
Walker C. 1983. Taxonomic concepts in the Endogonaceae: spore wall characteristics in species
descriptions. Mycotaxon 18: 443-455.
Walker C, Cuenca G, Sanchez F. 1998. Scutellospora spinosissima sp. nov. a newly described
Glomalean fungus from low nutrient communities in Venezuela. Annals of Botany 82:
721-725.
Walker C, Vestberg M. 1998. Synonymy amongst the arbuscular mycorrhizal fungi: Glomus claroideum
G. maculosum G. multisubstensum and G. fistulosum. Annals of Botany 82:601-624.
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MY COTA ZXON
Volume 105, pp. 89-93 July-September 2008
Arthonia hawksworthii sp. nov. (Ascomycota, Arthoniaceae)
on Dimelaena oreina from Turkey
MEHMET GOKHAN HALICI
'mghalici@erciyes.edu.tr
Biyoloji Boliimii, Fen Fakiiltesi, Erciyes Universitesi
38039 Kayseri, Turkey
Abstract — Arthonia hawksworthii is described on the thallus of Dimelaena oreina from
two localities in Turkey. It is most similar to A. gelidae, a lichenicolous species confined
to Placopsis gelida, but mainly differs from this species in that it has ascospores with a
gelatinous sheath that stain red in I-Lugol and an epithecium that is light to dark brown
with an olivaceous tinge in KOH. The new species is the first Arthonia species reported
on Dimelaena.
Key words —fungi, lichens, taxonomy
Introduction
The genus Arthonia Ach. includes c. 500 species, the majority of which are
lichenized (Hawksworth et al. 1995). The current concept of the genus is rather
broad, as it includes some species with and others without a K/I+ blue ring in
the tholus, and the interascal filaments are highly variable within the genus
(Coppins 1989, Grube & Matzer 1997). Grube & Matzer (1997) provided
useful taxonomical characters for Arthonia species, focusing primarily on
lichenicolous taxa.
Material and methods
The type of the new species is deposited in the herbarium at the Erciyes
University, Department of Biology, Kayseri. Specimens were examined with
an Olympus BH-2 research microscope fitted with Nomarski differential
interference contrast optics and a drawing tube. Sections were prepared by
hand and examined in I (Lugol’s iodine (MERCK 9261) and Meltzer’s iodine,
with [KI] and without [I] pre-treatment with 10% KOH), 10% KOH, and water.
Ascospore measurements were taken in water; the extreme values outside
the main range are seen in parentheses. The length/breadth (1/b) ratio of the
ascospores is provided in the same way.
90 ... Halici
The species
Arthonia hawksworthii Halici, sp. nov. FIGURE 1
MycoBank MB 511724
Fungus in thallo Dimelaenae oreinae incolens. Apothecia nigra, 0.1- 0.4 mm diam.
Epithecium 10-20 um altum, olivaceo-brunneum. Hymenium 50-65 um altum, hyalinum.
Hypothecium 10-15 ym altum, hyalinum vel pallide brunneum. Asci (23-)27-38 x 14-
18(-22) um, clavati, octospori. Ascosporae 1- septatae, hyalinae, halonatae, (10-)11.5-14
x 4-5.5(-6.5) um, l/b = (1.8-)2.3-3.0(-3.5).
Typus: Turkey, Yozgat, Cayiralan District, El¢i Village, 39°15’N, 35°39’E, alt. 1720 m, on
thallus of Dimelaena oreina on siliceous rocks, 14 August 2004, leg. M. G. Halici 0.1386
(herbarium of Erciyes University — holotypus).
EtyMoLocy: Named in honour of David L. Hawksworth, one of the pioneers of
lichenicolous fungi and a tutor of the author.
DESCRIPTION: Lichenicolous, on the thalli of Dimelaena oreina, causing slight
bleaching, weakly parasitic. AScoMATA apothecia, numerous, dispersed on the
surface of the host areoles, black, epruinose, superficial, plane at first, soon
becoming + convex, 0.1-0.4 mm diam, 2-4 per areole, lacking an exciple,
arthonioid. Epithecium light to dark olivaceous brown, 10-20 um tall, K-, N-;
hymenium colourless, 50-65 jm tall, I, + orange-red, KI,,+ slightly blue,
I etzer > HYpothecium colourless to pale brown, 10-15 um tall. HAMATHECIUM
of paraphysoids, abundant, septate, branched and anastomosed, 2-2.5 um
wide; markedly swollen apices, olivaceous brown, to 4-5.5 um wide. AscI
broadly clavate, short-stalked, bitunicate in structure, 8-spored, inner part
ee red, LSE ca without a So ag blue apical ring, (23-)27-38 x 14-18(-22)
uum. ASCOSPORES irregularly arranged in the asci, ellipsoid, hyaline, 1-septate,
rounded to somewhat broadly pointed at the apices, slightly constricted at the
septa, the upper cell slightly larger than the lower one, with 1-3 oil droplets at
times and a gelatinous sheath, the gelatinous sheath I,,+ red, (10-)11.5-14 x
4—5.5(-6.5) um (n = 36), I/b = (1.8-)2.3-3.0(-3.5), all measurements including
the closely adhering sheath. CoNIDIOMATA not observed.
ECOLOGY AND DISTRIBUTION: Commensalistic or weakly parasitic on the thalli
of Dimelaena oreina as slight bleaching occurs on the infected areoles of the
host thalli, especially in the later infection stage. The new species is known
from two localities in central and eastern Turkey. As the host species has a
wide distribution in the Northern Hemisphere, the species should be sought
for elsewhere.
OBSERVATIONS: Arthonia hawksworthii is most similar to A. gelidae R.Sant.,.
which was described by Santesson (1986) on the thallus of Placopsis gelida from
Sweden. Besides the different hosts, A. hawksworthii differs from A. gelidae in
that it has ascospores with a gelatinous sheath that stains red in I wgo, 20d has
an epithecium that is light to dark brown with an olivaceous tinge in KOH.
Arthonia hawksworthii sp. nov. (Turkey) ... 91
10 pm.
Fig. 1. Arthonia hawksworthii (holotype). A, mature asci. B, Six ascospore outlines showing the
gelatinous sheath, two of the ascospores including oil droplets. Scale = 10 um.
The lichenicolous species of Arthonia characterized by positive iodine reactions
of ascospore sheaths include A. pseudopegraphina Matzer, A. intermedia
Matzer, A. santessonii Matzer, A. diploiciae Calat. & Diederich, A. almquistii
Vain., A. amylospora Almq., A. intexta Almq. and A. obscurior Triebel (Grube
& Matzer 1997). The hymenium is also I, + dark blue and KI,,,,+ red in
A. gelidae (Santesson 1986), but I, ,,+ only slightly blue and KI,,,+ orange-red
in A. hawksworthii. A. epimela Norman, a lichenicolous species on Amandinea
punctata, has larger ascomata than A. hawksworthii [0.1-0.4 mm diam vs. 0.4-
0.6 mm diam], narrower ascospores [3.5-4.5 um vs. 4-5.5(-6.5) um] and a
brown hypothecium (Almquist 1880, Clauzade et al. 1989). A. epiphyscia Nyl.,
which is commensalistic or weakly parasitic on Physcia species, has a dark
reddish K+ olivaceous hypothecium (Coppins 1992). A. phaeophysciae Grube
& Matzer, a parasite causing considerable damage to Phaeophyscia thalli, has
asci with a KI + blue apical ring and a K+ greyish hypothecium (Grube &
Matzer 1997). A. caerulescens (Almq.) R.Sant., a lichenicolous species confined
to the apothecia of Lecanora varia, has shorter ascospores [10-12 um vs.
(10-)11.5-14 um] (Almquist 1880). The characters distinguishing the similar
species are provided in Table 1.
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Arthonia hawksworthii sp. nov. (Turkey) ... 93
ADDITIONAL SPECIMENS EXAMINED: Turkey, Kahramanmaras, Narli District, near the
highway to Gaziantep, 37°19’N, 37°08’E, alt. 600 m, on areoles of Dimelaena oreina
on siliceous rocks, 7 July 2004, leg. M. G. Halici 0.1387 (herbarium at the Erciyes
University).
Acknowledgements
Paul Diederich (Luxembourg) and Violeta Atienza (Spain) are thanked for reviewing this
paper. This study was carried out while I was in the Facultad de Farmacia, Universidad
Complutense de Madrid, under the direction of David L. Hawksworth, with a grant from
TUBITAK. Gokhan Hinisli prepared the figures. I also wish to express my gratitude to
Ahmet Aksoy and M. Yasar Dadand1 for their help in the fieldwork.
Literature cited
Almquist S. 1880. Monographia Arthoniarum Scandinaviae. Kongl Svenska Vetensk. Akad. Hand.
17: 1-69.
Clauzade G, Diederich P, Roux C. 1989. Nelikenigintaj fungoj likenlogaj. Bulletin de la Société
linnéenne de Provence, numéro spécial 1: 1-142.
Coppins BJ. 1989. Notes on the Arthoniaceae in the British Isles. Lichenologist 21: 195-216.
Coppins BJ. 1992. Arthonia Ach. (1806). In: Purvis OW, Coppins BJ, Hawksworth DL, James
PW, Moore DM (eds): The Lichen Flora of Great Britain and Ireland: 74-87. Natural History
Museum Publications, London.
Grube M, Matzer M. 1997. Taxonomic concepts of lichenicolous Arthonia species. Bibliotheca
Lichenologica 68: 1-17.
Hawksworth DL, Kirk PM, Sutton BC, Pegler DN. 1995. Ainsworth & Bisby’s dictionary of the
fungi. 8"" edn. Wallingford.
Santesson R. 1986. Fungi lichenicoli exsiccatae. Fasc. 3-4 (Nos 51-100). Thunbergia 3: 1-18.
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MYCOTAXON
Volume 105, pp. 95-104 July-September 2008
Corticolous lichens of the city of Giresun
with descriptions of four species new to Turkey
KENAN YAZICI** & ANDRE APTROOT?
*kcagri_1997@yahoo.com
'Karadeniz Technical University, Faculty of Sciences and Letters
Biology Department, Trabzon, Turkey
andreaptroot@wanadoo.nl
*Adviesbureau voor Bryologie en Lichenologie
G.v.d. Veenstraat 107, NL-3762 XK Soest, The Netherlands
Abstract — Corticolous lichens were studied in the city of Giresun (Turkey). The
survey yielded 54 species, all Ascomycota, among which Anisomeridium polypori,
Lecanora compallens, Lepraria umbricola and Schismatomma decolorans are new to
Turkey. Lecanora compallens is new to Asia while Anisomeridium polypori and Lepraria
umbricola are reported for the second time from Asia. Data and accompanying species
are given for all taxa while short descriptions are also presented for the four species that
are new to Turkey.
Keywords — biodiversity, flora
Introduction
Studies on the lichen flora of Turkey are not as extensive as elsewhere. Last
year, many new lichen taxa were reported for Turkey (Dogru & Giiven¢
2007, Yazici & Aptroot 2007, Yazici et al. 2007a,b). Although there was some
previous lichenological research in the Black Sea Coast city of Giresun (Aslan
& Yazici 2006, Duman & Yurdakulol 2007, Kinalioglu 2006), several regions in
Giresun remained still unexplored. The present paper provides information on
corticolous lichens in the city center of Giresun, Turkey.
Materials and methods
The corticolous lichen samples were collected from deciduous trees from
01-30 October 2007 in the city of Giresun (Turkey). The study area is situated
at 40°54N, 38°23E. Air-dried lichen samples were observed and studied using
stereo and light microscopes. Secondary metabolites were identified by the
usual spot tests (Poelt 1974, Purvis et al. 1992, Herk & Aptroot 1999, Wirth
1995, Zedda 2000).
96 ...Yazici & Aptroot
Results
A lichen survey of the city of Giresun yielded 54 species, all Ascomycota, among
which Anisomeridium polypori, Lecanora compallens, Lepraria umbricola and
Schismatomma decolorans are new to Turkey. Lecanora compallens is new to
Asia, while Anisomeridium polypori and Lepraria umbricola are reported
here for the second time from Asia. Additionally, Caloplaca flavocitrina and
Phaeophyscia chloantha represent second reports for Turkey. Collection data
and information accompanying species are given for all lichen taxa, and short
descriptions are also presented for the four species new to Turkey. The lichen
taxa are listed alphabetically.
Amandinea punctata (Hoftm.) Coppins & Scheid.
City center and Giresun Castle, on Fraxinus sp., Albizia julibrissin, main roadside on
Robinia pseudoacacia, accompanying species: Caloplaca obscurella, Candelariella reflexa,
Hyperphyscia adglutinata, Lecanora compallens, Phaeophyscia orbicularis and Rinodina
gennarii. Yazici 1304.
Anisomeridium polypori (Ellis & Everh.) M.E. Barr
Thallus mostly inconspicuous, effuse, grey or pale grey to green. Perithecia
0.15-0.25 mm diam., subconical to + globose, initially immersed; upper wall
with a slightly converted involucrellum. Asci 5-75(-85) x 11-16 um, cylindric-
clavate. Ascospores (13-)15-20(-22) x (3.5-)4.5-5 um, 1-3 septate, clavate-
fusiform, Pycnidia black, of two types, first one: 100-150 um diam., + sessile,
conical, colourless, with ellipsoid or ovoid macroconidia, (3-)3.5-4.5(-5) x 1.8-
2 um, extruded as a white cirrus, in which the conidia are bound by a gelatinous
matrix; second one: 50-100 um diam., + immersed, globose, with rod-shaped
to narrowly ellipsoid microconidia, 2-3 x 1-1.3 um. Photobiont Trentepohilia.
Known throughout Europe, Australia, Canada, China, Costa Rica, Japan,
North America, Russia, Thailand. A second report for Asia.
Giresun: Center; Giresun Castle, 120 m, 40°54’57” N, 38°23’24” E, on Fraxinus sp.,
unpolluted area, accompanying species: Gyalecta truncigena. 10 October 2007. Yazici
1318.
ECOLOGY OF THE SURVEY AREA-The area has a typical oceanic climate with a
mild and high precipitation prevailing along the coast. It is at 120 m elevation
and at 150 m distance from the Black Sea. Annual temperature is 14.2°C in the
city of Giresun while annual precipitation is 1305 mm. On average there are 184
rainy days, 6 snowy days and 11 snow-covered days. Winters are more intense
with longer snow cover, while the summers are cool. Rainy days occur in every -
season, the most rainy days however in autumn and winter. The well-lit site,
which is occasionally exposed to strong wind in winter, is rich compared to the
forest. It has tree communities with Albizia julibrissin, Robinia pseudoacacia,
Salix sp. and Platanus orientalis.
Corticolous lichens of Giresun (Turkey) ... 97
REMARKS—Anisomeridium polypori is a cosmopolitan species frequent
suboceanic and mild-temperate region. Generally it especially grows on
rough bark of Sambucus, Populus, and Ulmus by rivers and brooks in humid
woodland, rarely on shaded rock, and is sometimes common in polluted
areas. Anisomeridium polypori is very similar to A. biforme, but it has a less
conspicuous thallus, smaller perithecia with a thinner upper wall, and longer
and narrower ascospores with often 1-3 septa, but is especially characterized by
the long-ostiolate pycnidia with agglutinating conidia.
Arthonia dispersa (Schrad.) Nyl.
City center, on Acer sp., Fraxinus sp., Ficus sp.; Giresun Castle, on Albizia julibrissin,
Ficus sp., Fraxinus sp., and Salix sp., accompanying species: Naetrocymbe punctiformis.
Yazici 1323.
Arthonia radiata (Pers.) Ach.
Giresun Castle, 120 m, on Robinia pseudoacacia and Fraxinus sp. Yazici 1322.
Bacidia laurocerasi (Duby) Zahlbr.
City center, on Ligustrum sp., Betula sp.; Giresun Castle, 120 m, on Fraxinus sp. Yazici
Keay
Caloplaca cerina (Ehrh. ex Hedw.) Th. Fr.
City center and main roadside on Robinia pseudoacacia, accompanying species: Physcia
adscendens, P. tenella and Xanthoria parietina. Yazici 1327.
Caloplaca cerinella (Nyl.) Flagey
City center, on Acer sp., accompanying species: Hyperphyscia adglutinata, Lecania
fuscella and Lecidella achristotera. Yazici 1301.
Caloplaca citrina (Hoffm.) Th. Fr.
City center, park, on Salix sp., Platanus orientalis, Robinia pseudoacacia, and Ligustrum
sp., main roadside on Fraxinus sp. and Robinia pseudoacacia; Giresun Castle, on Albizia
julibrissin, accompanying species: Hyperphyscia adglutinata, Phaeophyscia orbicularis,
Physcia tenella, P. adscendens and Xanthoria parietina. Yazici 1328.
Caloplaca flavocitrina (Nyl.) H. Olivier
City center and main roadside, on Robinia pseudoacacia, accompanying species:
Candelariella reflexa, Lecanora dispersa and Phaeophyscia orbicularis. Found for the
second time in Turkey. Yazici 1305.
Caloplaca holocarpa (Hoffm.) A.E. Wade
City center, on Acer sp. and Robinia pseudoacacia, accompanying species: Caloplaca
cerina, Hyperphyscia adglutinata, Lecania cyrtella, Lecanora chlarotera, L. dispersa,
Lecidella elaeochroma, Physcia adscendens and P. tenella. Yazici 1307.
Caloplaca obscurella (J. Lahm ex K6rb.) Th. Fr.
City center, on Robinia pseudoacacia, accompanying species: Amandinea punctata,
Hyperphyscia adglutinata, Phaeophyscia orbicularis and Rinodina gennarii. Yazici 1303.
98 ...Yazici & Aptroot
Candelaria concolor (Dicks.) Stein
City center and main roadside, on Fraxinus sp., accompanying species: Hyperphyscia
adglutinata, Lecanora dispersa, Physcia adscendens, P. tenella and Xanthoria parietina.
Yazici 1329.
Candelariella aurella (Hoffm.) Zahlbr.
City center and main roadside, on Robinia pseudoacacia, accompanying species:
Lecanora dispersa. Yazici 1330.
Candelariella reflexa (Nyl.) Lettau
City center, on Robinia pseudoacacia and Fraxinus sp., accompanying species: Amandinea
punctata, Caloplaca flavocitrina, Lecania cyrtella, Lecanora compallens, L. dispersa and
Phaeophyscia orbicularis. Yazici 1302.
Chrysothrix candelaris (L.) J.R. Laundon
Giresun Castle, 120 m, on Albizia julibrissin. Yazici 1331.
Evernia prunastri (L.) Ach.
Giresun Castle, 120m, on Albizia julibrissin, accompanying species: Flavoparmelia
caperata and Pseudevernia furfuracea. Yazici 1332.
Flavoparmelia caperata (L.) Hale
City center, 5 m, on Pyrus sp., main roadside; Giresun Castle, 120 m, on Robinia
pseudoacacia and Albizia julibrissin, accompanying species: Punctelia subrudecta,
Parmelia sulcata, Evernia prunastri and Melanelixia fuliginosa. Yazici 1334.
Graphis scripta (L.) Ach.
Giresun Castle, 120 m, on Albizia julibrissin. Yazici 1321.
Gyalecta truncigena (Ach.) Hepp
Giresun Castle, on Fraxinus sp. accompanying species: Anisomeridium polypori. Yazici
PS1S:
Hyperphyscia adglutinata (Flérke) H. Mayrhofer & Poelt
City center and along the main roadside 5-20 m, on Tilia sp., Acer sp., Salix sp., Robinia
pseudoacacia, Ligustrum sp., Cercis siliquastrum and Fraxinus sp., also Giresun Castle
120 m, on Albizia julibrissin and Platanus orientalis, accompanying species: Amandinea
punctata, Caloplaca cerinella, C. holocarpa, C. obscurella, Lecania fuscella, Lecidella
achristotera, Phaeophyscia chloantha, P. orbicularis and Rinodina gennarii. Yazici 1303.
Hypogymnia physodes (L.) Nyl.
Giresun Castle, 120 m, on Albizia julibrissin, accompanying species: Hypogymnia
tubulosa, Yazici 1333.
Hypogymnia tubulosa (Schaer.) Hav.
Giresun Castle, 120 m, on Albizia julibrissin, accompanying species: Hypogymnia
physodes. Yazici 1335.
Corticolous lichens of Giresun (Turkey) ... 99
Lecania cyrtella (Ach.) Th. Fr.
City center, 40 m, on Acer sp., main roadside, on Robinia pseudoacacia, accompanying
species: Candelariella reflexa and Lecanora umbrina. Yazici 1316.
Lecania fuscella (Schaer.) A. Massal.
City center, 40 m, on Acer sp., main roadside, on Robinia pseudoacacia, accompanying
species: Caloplaca holocarpa, C. cerinella, Hyperphyscia adglutinata, Lecidella achristotera,
Physcia adscendens and Lecanora dispersa. Yazici 1307.
Lecania naegelii (Hepp) Diederich & Van den Boom
City center, 40 m, on Acer sp., accompanying species: Lecania cyrtella. Yazici 1336.
Lecanora argentata (Ach.) Malme
City center, 5 m, on Salix sp.; Giresun Castle, 120 m, on Albizia julibrissin, Castanea
sp., and Acer sp., main roadside, 5 m, on Ficus sp., accompanying species: Lecidella
achristotera. Yazici 1310.
Lecanora campestris (Schaer.) Hue
City center, 5 m, on Ficus sp. Yazici 1337.
Lecanora chlarotera Ny.
City center, 5m, on Salix sp. and Acer sp., accompanying species: Caloplaca holocarpa.
Vazicw lle
Lecanora compallens Herk & Aptroot
Thallus corticolous, usually 1-3 cm diam., continuous; margin whitish grey,
immersed and shiny, usually with low, glaucous grey warts of 0.1-0.2 mm
diam., without hypothallus; medulla below soredia 0.1-0.3 mm thick, white.
Soralia always present, usually covering most of the thallus, except for a
marginal zone of c. 1 mm., 0.1-0.3 mm punctiform openings, soon uprising
and aggregating into irregular patches. Soredia granular, in a dense mass, 15-
30 in diam., yellowish to slightly mint-green, contrasting in colour with the
thallus. Apothecia and pycnidia unknown. Thallus C- , P- , K-, UV-; soredia
C- , P-, K+ yellowish to yellowish brown usnic acid and zeorin found with TLC
(Herk & Aptroot 1999).
Lecanora compallens, previously known from Western Europe, most
commonly occurs on medium-aged trees, mainly on the flat surfaces of the
ridges, and also on acid as well as neutral bark. Usually on the west side of
exposed wayside trees, especially Quercus, Fraxinus, Populus, Salix, Sorbus,
Tilia, Ulmus and Sambucus.
Known from Belgium, Denmark, England, France, Germany, Lithuania,
Luxembourg, Poland, the Netherlands, and Ukraine. New to Asia.
Giresun: Center: Giresun Castle, 40°545°7” N, 38°23’24” E, on Albizia julibrissin, 120 m,
accompanying species: Amandinea punctata, Phaeophyscia orbicularis and Candelariella
reflexa. 10 October 2007. Yazici 1302.
100 ... Yazici & Aptroot
ECOLOGY OF THE SURVEY AREA-See Anisomeridium polypori.
REMARKS — Lecanora compallens is very similar to L. expallens and L.
barkmaniana, but L. compallens is distinguished from L. expallens in always
having a non-sorediate thallus margin of 1 mm with low glaucous grey
corticate warts. Furthermore, L. expallens has usually excavate soralia and a
more yellowish, never mint-green, colour, and sometimes a black hypothallus
is present. L. expallens can be easily separated by the C +orange reaction due
to the presence of thiophanic acid. L. barkmaniana and L. compallens have
microscopically identical soredia, but L. compallens differs by the yellowish
tinge of the soredia and by a much thinner, often mostly immersed thallus, and
L. barkmaniana reacts Pd+ yellow.
Lecanora dispersa (Pers.) Sommerf.
City center, on Fraxinus sp., Pyrus sp., and Robinia pseudoacacia, main roadside, on Acer
sp., Fraxinus sp., Salix sp., and Robinia pseudoacacia, accompanying species: Caloplaca
flavocitrina, C. holocarpa, Candelariella aurella, C. reflexa, Lecania fuscella, Physcia
adscendens, Phaeophyscia orbicularis and Rinodina gennarii. Yazici 1309.
Lecanora umbrina (Ach.) A. Massal.
City center and main roadside, 2 m, on Robinia pseudoacacia and Acer sp., accompanying
species: Lecania cyrtella. Yazici 1316.
Lecidella achristotera (Nyl.) Hertel & Leuckert
City center and main roadside, on Acer sp., Ficus sp., Fraxinus sp., Pyrus sp.; Giresun
Castle, 120 m, on Robinia pseudoacacia, accompanying species: Caloplaca cerinella,
Hyperphyscia adglutinata, Lecania fuscella, Lecanora argentata and Lecidella elaeochroma.
Yazici 1301.
Lecidella elaeochroma (Ach.) M. Choisy
City center, 10 m, on Acer sp., Ligustrum sp., and Prunus sp., main roadside, 2 m, on Ficus
sp. and Robinia pseudoacacia; Giresun Castle, 120 m, on Albizia julibrissin, Laurus sp.,
and Robinia pseudoacacia, accompanying species: Caloplaca holocarpa and Xanthoria
parietina. Yazici 1338.
Lepraria incana (L.) Ach.
Giresun Castle, 120 m, on Albizia julibrissin. Yazici 1339.
Lepraria umbricola Tonsberg
Thallus leprose, powdery appearance, diffuse, deep greenish or whitish greyish,
sometimes weft—like, usually thin, rarely thick, cortex absent, lacking marginal
lobes and without medulla, protected from direct rain in shaded sites; soredia _
abundant to scattered, forming small granules, granules to 0.1 mm diam and
short projecting hyphae frequent. Thamnolic acid present, thallus K + deep
yellow, C-, KC-, P + deep orange.
Corticolous lichens of Giresun (Turkey) ... 101
Lepraria umbricola grows on bark, rock, mosses, soil, mostly acidic substrata;
shaded, sheltered, humid, sometimes extremely shaded.
Known throughout Europe, Africa, Australia, Canada, Colombia, North
America, Pacific Northwest, Russia, and even (probably incorrectly) reported
from Thailand. Found for the second time in Asia.
Giresun: Center: Giresun Castle, 40°545’7” N, 38°23'24” E, on Albizia julibrissin, 120 m.
10 October 2007. Yazici 1308.
ECOLOGY OF THE SURVEY AREA—See Anisomeridium polypori.
REMARKS-Lepraria umbricola is an oceanic to warm-—temperate species, found
on sheltered mossy siliceous rocks and terricolous mosses, sometimes on basal
parts of old trunks on shaded sandy soil. Lepraria umbricola is somewhat similar
to L. incana and L. eburnea, but L. incana has a more conspicuous medulla.
Additionally L. incana has divaricatic acid and zeorine while L. umbricola has
thamnolic acid.
Lepraria vouauxii (Hue) R.C. Harris
City center, 20 m, on Tilia sp., main roadside, 2 m, on Magnolia sp. Yazici 1313.
Melanelixia fuliginosa (Fr. ex Duby) O. Blanco et al.
Giresun Castle, 120 m, on Albizia julibrissin, accompanying species: Flavoparmelia
caperata and Parmelia sulcata. Yazici 1340.
Melanelixia subaurifera (Nyl.) O. Blanco et al.
Giresun Castle, 120 m, on Albizia julibrissin. Yazici 1319.
Naetrocymbe punctiformis (Pers.) R.C. Harris
Giresun Castle, 120 m, on Albizia julibrissin, Fraxinus sp., and Prunus sp., accompanying
species: Arthonia dispersa. Yazici 1320.
Opegrapha atra Pers.
City center, 50 m, on Corylus sp., Quercus sp.; Giresun Castle, 120 m, on Acer sp., Corylus
sp., and Fraxinus sp. Yazici 1325.
Opegrapha varia Pers.
City center, 2 m, on Acer sp.; Giresun Castle; 120 m, on Acer sp. Yazici 1324.
Opegrapha vulgata (Ach.) Ach.
City center, 10 m, on Morus sp. Yazici 1326.
Parmelia sulcata Taylor
City center, main roadside, 2 m, on Fraxinus sp.; Giresun Castle, 120 m, on Albizia
julibrissin, accompanying species: Flavoparmelia caperata and Melanelixia fuliginosa.
Yazici 1341.
Pertusaria amara (Ach.) Nyl.
Giresun Castle, 120 m, on Albizia julibrissin. Yazici 1342.
102 ...Yazici & Aptroot
Phaeophyscia chloantha (Ach.) Moberg
City center, main roadside, 2 m, on Magnolia sp. and Robinia pseudoacacia, Giresun
Castle, 120 m, on Albizia julibrissin, accompanying species: Hyperphyscia adglutinata
and Phaeophyscia orbicularis. Found for the second time in Turkey. Yazici 1311.
Phaeophyscia orbicularis (Neck.) Moberg
City center, 5-50 m, on Acer sp., Ligustrum sp., main roadside, 2 m, on Robinia
pseudoacacia, Acer sp., Fraxinus sp., and Populus sp.; Giresun castle, 120 m, on Albizia
julibrissin, accompanying species: Amandinea punctata, Caloplaca flavocitrina, C.
obscurella, Candelariella reflexa, Hyperphyscia adglutinata, Lecanora compallens, L.
dispersa, Phaeophyscia chloantha, Physcia tenella, Rinodina gennarii and Xanthoria
parietina. Yazici 1302.
Physcia adscendens (Th. Fr.) H. Olivier
City center, 5—50 m, on Acer sp., Fraxinus sp., Ligustrum sp., Populus sp, and Robinia
pseudoacacia; Giresun Castle, 120 m, on Albizia julibrissin, Fraxinus sp., Acer sp.,
accompanying species: Caloplaca cerina, C. holocarpa, Phaeophyscia orbicularis and
Physcia tenella. Yazici 1307.
Physcia tenella (Scop.) DC.
City center, 10 m, on Ligustrum sp., main roadside, 2m, on Robinia pseudoacacia and
Populus sp., accompanying species: Caloplaca cerina and Physcia adscendens. Yazici
1343.
Punctelia subrudecta (Nyl.) Krog
City center, 2 m, on Pyrus sp., accompanying species: Flavoparmelia caperata. Yazici
1344.
Pseudevernia furfuracea (L.) Zopf
Giresun Castle, 120 m, on Albizia julibrissin, accompanying species: Evernia prunastri
and Flavoparmelia caperata. Yazici 1345.
Ramalina farinacea (L.) Ach.
Giresun Castle, 120 m, on Albizia julibrissin. Yazici 1346.
Rinodina gennarii Bag.
City center and main roadside, 2 m, on Robinia pseudoacacia, accompanying species:
Amandinea punctata, Caloplaca obscurella, Hyperphyscia adglutinata, Lecanora dispersa
and Phaeophyscia orbicularis. Yazici 1306.
Schismatomma decolorans (Turner & Borrer ex Sm.) Clauzade & Vézda
Thallus crustose, sorediate, thin, to 3 cm in diam., attached to the substrate
with the whole lower surface or with raised, free margins lilac to pinkish grey, —
mauve-grey to brown, developing concolorous or slightly paler sorediate
patches which usually cover the whole surface of mature specimens, giving
a leprose appearance, becoming grey to brownish grey in the herbarium, +
rimose-—cracked; a grey-black prothalius often + present; with entire margins
Corticolous lichens of Giresun (Turkey) ... 103
and continuous or more ore less fissured and divided into areoles, flat and
mainly parallel to the substrate; upper side clearly different from the underside,
not much swollen when wet and usually not wrinkled when dry; photobiont
Trentepohlia, algal cells and hyphae surrounded by air. Soredia puntiform,
0.2-0.8(-1) mm daim., + concolorous discrete or sparingly confluent, often
covering relatively large areas, mauve-grey to pale lilac-grey, occasionaly with
an ochry tinge. Apothecia and pycnidia unknown. Thallus K-, KC-, C-, P-,
UV-.
Known throughout Europe (e.g. eastern part of the Italian peninsula,
England, France, Germany, Northern Ireland, Spain, Sweden, The Netherlands),
Algeria, Cyprus, Iran, North America, Syria, Tanzania. Reported for the second
time from Asia.
Giresun: Center; Giresun Castle, 40°545°7” N, 38°23°24” E, on Albizia julibrissin, 120 m,
10 October 2007. Yazici 1314.
ECOLOGY OF THE SURVEY AREA-See Anisomeridium polypori.
REMARKS-Schismatomma decolorans is a mild—temperate and suboceanic
species, most commonly growing in sheltered and shaded recesses of old rough-
barked deciduous trees in wayside and woodland sites, rarely on dry vertical
rocks or walls, exposed mineral-rich rock stones and steeply inclined faces
in lichen—poor communities, tolerant of relatively high levels of agricultural
chemicals and fertilizers.
Xanthoria parietina (L.) Th. Fr.
City center, 2-50 m and main roadside, 2 m, on Ligustrum sp., Acer sp., Elaeagnus sp.,
Ficus sp., Fraxinus sp., Platanus orientalis, Pyrus sp., Robinia pseudoacacia, and Salix
sp., accompanying species: Caloplaca cerina, Candelaria concolor, Lecidella elaeochroma,
Physcia adscendens and Phaeophyscia orbicularis. Yazici 1312.
Discussion
The population of Giresun city at the Eastern Black Sea coast is approximately
100,000, according to the census in 2007. The average elevation of the city center
is 50 m. Giresun Castle, which is situated in the north of the city, lies at 120 m
elevation and at 150 m distance from the Black Sea, in an unpolluted area. Until
2003, sulphur dioxide measurements from the area are not reliable, and these
measurements were not performed in 2003—2006. From January 2007 onwards,
more reliable measurements have been made. According to these, the annual
average sulphur dioxide concentration in the atmosphere is approximately
50ug/m?in 2007 (Turkiye Cumhuriyeti Cevre ve Orman Bakanligi 2007).
In total 33 lichen species, including the four new to Turkey, were found in
Giresun Castle, while 30 species were present along the main roads; 36 taxa
in the other parts of the city center. All species were found on 20 different
104 ...Yazici & Aptroot
deciduous tree species, such as Acer sp., Albizia julibrissin, Ficus sp., Fraxinus
sp., Ligustrum sp., Robinia pseudoacacia and Salix sp.: 26 species were found
on Albizia julibrissin, 23 on Robinia pseudoacacia, 18 on Acer sp. and 18 on
Fraxinus sp. The area Giresun Castle is apparently an unpolluted area since its
33 lichen taxa include sensitive species such as Evernia prunastri, Pseudevernia
furfuracea, Ramalina farinacea, Hypogymnia physodes and H. tubulosa. On the
other hand, it is remarkable that still 30 lichen species have been found along
the main roads, although these sites are exposed to heavy metals from motor
vehicles and other forms of pollution.
Acknowledgements
We are grateful to Professor Orvo Vitikainen and Dr. Leo Spier for linguistic revision
and helpful comments on an earlier draft of this manuscript. We also acknowledge the
assistance of Dr. Piercey-Normore and Dr. Guttova here.
Literature cited
Aslan A, Yazici K. 2006: Contribution to the lichen flora of Giresun province of Turkey. Acta Bot.
Hung. 48: 231-245.
Dogru Z, Giiveng S. 2007: Lichenized and lichenicolous fungi from Bursa province new to Turkey.
Mycotaxon 102: 389-394.
Duman DC, Yurdakulol E. 2007: Lichen Records from Saricgicek Mountain in Southern Giresun
Province, Turkey. Turk. J. Bot. 31: 357-365.
Herk CM van, Aptroot A. 1999: Lecanora compallens and L. sinuosa, two new overlooked corticolous
lichen species from Western Europe. Lichenologist 31: 543-553.
Kinalioglu K. 2005: Lichens of Giresun District Giresun Province, Turkey. Turk. J. Bot 29:
417-423.
Poelt J. 1974: Bestimmungsschliissel Europaischer Flechten. J.Cramer, Lehre.
Purvis OW, Coppins BJ, Hawksworth DL, James PW, Moore DM. 1992: The Lichen Flora of Great
Britain and Ireland. Natural History Museum & British Lichen Society, London.
Turkiye Cumhuriyeti Cevre ve Orman Bakanligi. Tiirkiye Hava Izleme Agi, Hava Izleme Raporu,
2007.
Wirth V. 1995: Die Flechten Baden-Wiirttembergs. Teil 1-2. Ulmer, Stuttgart.
Zedda L. 2000: The lichen genera Lepraria and Leproloma in Sardinia (Italy). Cryptogamie Mycol.
21: 249-267:
Yazici K, Aptroot A. 2007: Five lichens new to Turkey. Mycotaxon 100: 21-26.
Yazic1 K, Aptroot A, Aslan A. 2007a: Six lichenized and non-lichenized fungi new to Turkey.
Mycotaxon 102: 307-313.
Yazic1 K, Aptroot A, Aslan A. 2007b: Lichen biota of Zonguldak, Turkey. Mycotaxon 102:
257-260.
MYCOTAXON
Volume 105, pp. 105-110 July-September 2008
Balaniopsis triangularis sp. nov. from indoor environments
DeE-WEI Li’, BRYCE KENDRICK’,
DAVID SPERO? & CLAIRE MACDONALD4
dewei.li@po.state.ct.us
' The Connecticut Agricultural Experiment Station, Valley Laboratory
153 Cook Hill Road, Windsor, CT 06095
bryce@mycolog.com
’ 8727 Lochside Drive, Sidney, BC, V8L 1M8, Canada
3 Natural Link Mold Lab
4900 Mill Street, Suite 3, Reno, NV 89502
*SanAir Technologies Laboratory
1551 Oakbridge Dr, Suite B, Powhatan, VA 23139
Abstract — A new species of Balaniopsis is described and illustrated from specimens
collected during indoor mold inspections in North America.
Key Words — hyphomycete, mitosporic fungi, taxonomy, type
Introduction
In recent years, unidentified triangular spores that appeared to be conidia
were recorded by the authors on tape lifts and in spore traps. Occasionally,
conidiogenous cells and parts of conidiophore stipes were also seen. On rare
occasions, the orientation of the spores relative to the conidiogenous cells and
conidiophores was seen. Although the organism has not been successfully
grown in culture, a specimen recently collected from conifer wood over a crawl
space in a building in Vancouver, Canada, allowed the authors to characterize
the species as Balaniopsis triangularis anam. sp. nov..
Materials and methods
Conidiophores and conidia of the fungus were mounted in lacto-fuchsin
(0.1 g acid fuchsin, 100 ml 85% lactic acid) (Carmichael 1955). Microscopic
* Corresponding author
106... Li & al.
observations were made using Nomarski differential interference contrast
optics. Herbarium acronyms follow Index Herbariorum (Holmgren &
Holmgren 1998).
Results
Balaniopsis triangularis D.W. Li & W.B. Kendr., anam. sp. nov. Figures 1-13
MycoBAnkK MB 511446.
Conidiophora macronemata, determinata, erecta, non-ramosa vel ramosa, recta vel
flexuosa, lutea, septata, interdum >200 um longa et 0.7-2 um crassa apud basis, 2.9-3.4
um apud apicem. Cellulae conidiogenae polyblasticae, laeves, cupulatae vel obconicae.
Conidia unicellularia, triangularia, non catenata, laevia, lutea, 6.0-7.4 x 6.4-8.6 x 3.0-
3.6 um. Teleomorphosis ignota.
Holotypus BPI 878719, Isotypus DAOM 239847 superficie in ligno domi, Vancouver,
Canada leg. S. Campbell,. 28 iv. 2008.
EryMoLocy: referring to the triangular shape of the conidia.
CONIDIOPHORES solitary, determinate, erect, unbranched or branched, straight
or flexuous, yellowish, smooth, septate, thin-walled when young, becoming
thick-walled at maturity, sometimes exceeding 200 um in length, 0.7-2 um
wide at base, widening to 2.9-3.4 um at the apex.
CONIDIOGENOUS CELLS terminal, subapical or lateral, obovoid first, becoming
cupulate or obconic (champagne flute-shaped) after conidia have been
released, due to partial collapse of apical portion, smooth, yellowish, terminal,
solitary, (6.8—) 9.3-14.7 (-16.6) (mean = 12 + 2.7, n=17) long, (5.6-) 6.3-7.9
(-8.6) (mean = 7.1 + 0.8 ttm, n=17) wide at the widest point, narrowing to the
proximal end (2.6-) 2.7-3.3 (-3.8) (mean = 3.0 + 0.3 um, n=17), subapical
or lateral in whorls of 3-5, (7.8—) 8.3-10.5 (-12.6) (mean = 9.4 + 1.1, n = 19)
x (5.1-) 6.6-8.4 (-8.9) (mean = 7.5 + 0.9, n = 19). Developing 6-8 conidia
polyblastically and perhaps synchronously in the apical area.
Conip1A 1-celled (amerosporous), cuneiform (wedge-shaped) when young,
triangular at maturity, yellowish, smooth-walled, later becoming thick-walled,
(5.6-) 6.0-7.4 (-8.8) (mean = 6.7 + 0.7, n = 31) x (5.5-) 6.4-8.6 (-9.6) um (mean
= 7.5 + 1.1 um, n = 31) x (2.7-) 3.0-3.6 (-4.2) um (mean = 3.3 + 0.3 um, n
= 31) um, side wall about 1 um thick, at the three corners 2-2.4 um thick.
Conidial secession is rhexolytic. The remnants of separating cell walls persist
at the base of some conidia (Figure 7). The corners of the released conidia may
appear rounded or, more commonly, truncate. The truncate condition probably
indicates some minor collapse of the outer wall subsequent to drying.
TELEOMORPH unknown.
KNOWN GEOGRAPHICAL DISTRIBUTION: British Columbia, Canada; California
and New York, USA.
—_—
Balaniopsis triangularis sp. nov. ... 107
20.00 um
F880 um
Figures 1-5. Balaniopsis triangularis. 1. Branched conidiophore, conidiogenous cells, and
conidia. 2. Conidiophore with terminal and lateral conidiogenous cells, and conidia. 3, 4.
Terminal conidiogenous cells. 5. Terminal and lateral conidiogenous cells. 5. Conidiogenous
cells. Scale bars: 1 = 20 um, 2, 5 = 10 um, 3-4 = 5 um.
HABITAT: saprotrophic on damp wood and plywood.
SPECIMENS EXAMINED: CANADA, British Columbia, Vancouver, Marine Drive, crawl
space of building, 28 April 2008, Shannon Campbell, holotype (BPI 878719), isotype
(DAOM 239847); UNITED STATES, New York, Livingston, 7 March 2007, anonymous,
(BPI 878411).
108 ::.. Mer Sal:
5.00 um
Figures 6-13. Balaniopsis triangularis. 6. Terminal conidiogenous cells. 7-8. Lateral
conidiogenous cells. 9. Lateral conidiogenous cells bearing conidia in face- and side-view
10-12. Conidia in face-view. 13. Conidia in face- and side-view.
Scale bars: 6, 8-12 = 5 um, 7, 13 = 10 um.
Discussion
The genus Balaniopsis P.M. Kirk was proposed by Kirk (1985) with a single
species, Balaniopsis africana (Kiffer) P.M. Kirk (= Balanium africanum Kiffer).
His establishment of the new genus was based on three major characters:
“+.
Balaniopsis triangularis sp. nov. ... 109
1) rhexolytic conidial secession, 2) percurrent extension, and 3) conidia in
short, branched chains on unbranched conidiophores. However, Whitton et
al. (2002) emended the generic description, reinterpreting the conidiogenous
cells and adding a second species. There were significant differences between
Kiffer’s specimen referred to as the holotype by Kirk (1985) and Kirk’s generic
and specific descriptions, which were based on the specimen he collected
in Kenya. The key characters of the revised circumscription of Balaniopsis
include rhexolytic conidial secession, sometimes with percurrent extension
of the conidiogenous cell, and unbranched or branched conidiophores. The
description of conidia as occurring in short branched chains was not included
in the revised circumscription because the entity defined by Kirk (1985) as a
basal conidium was interpreted by Ellis (1976) and Whitton et al. (2002) as a
conidiogenous cell, which implied that the conidia were produced singly and
probably synchronously.
Whitton et al. (2002) clarified that the holotype and Kirk’s specimen are
two different species. Balanium africanum, described and illustrated by Kiffer
(1973), has a light-dark-light pattern of pigmentation of the conidiophore cells,
triangular conidia (9-12 x 7-10 um), and longer separating cells, while Kirk’s
specimen has concolourous conidiophores, broadly obovoid conidia (12.8-14.5
x 9.5-11 um), and much shorter separating cells. The name Balaniopsis africana
must be applied to Kiffer’s collection. The specimen collected by Kirk is not
conspecific with that of Kiffer. Thus, Whitton et al. (2002) re-described it under
a new specific name, Balaniopsis kirkii Whitton et al. Another new species,
Balaniopsis dendroidea Whitton et al., with globose conidia and branched
conidiophores, was described by Whitton et al. (2002). Balaniopsis triangularis
is different from the other three species in its much smaller, triangular conidia
(6.0-7.4 x 6.4-8.6 um), obovoid or obconic conidiogenous cells, and reduced
separating cell.
Balaniopsis triangularis is in some ways similar to Stellomyces kendrickti Kesh.
Prasad & Bhat (Keshava Prasad & Bhat 2002). However, although the conidia
of S. kendrickii are often triangular, they have an attenuated base and arise from
very narrow extensions of the conidiogenous cell. In addition, in Stellomyces,
one conidiogenous cell may give rise to several clusters of conidia (which may
be terminal or intercalary) and that conidiogenous cell may even be branched,
with more than one cluster at the end of each branch. In B. triangularis, the
conidiogenous cell is not branched; each conidiogenous cell gives rise to only
one cluster of conidia which is always terminal; and the conidia do not arise on
fine outgrowths like those seen in Stellomyces.
The holotype was collected from a wooden structure in crawl space of a
building and could not be isolated in pure culture. It would appear that the
material had been in situ for so long that the mycelium and spores had died.
P10... Sbreeal
Although rare, this fungus occurs with significant frequency on building
materials, where its presence needs to be acknowledged and recorded. It is our
opinion that it is now necessary and appropriate to make available a binomial
for this organism. |
Key to species of Balaniopsis
1. Conidiophores typically dichotomously branched; conidia brown to pale brown,
mioreor less:spherical? 3:5 Spimidiameterssess..miete ne eee B. dendroidea
l.:Conidiophores ih branched itt Ws aeloe rota) Set. cht tn oti Oe eee aneS 2,
2. Conidia broadly obovoid, dark brown, 12.8-14.5 x 9.5-llum ............ B. kirkii
2*Gonidiamorcom less trials Ulqiempancy. Gee. tsee eeenet eet ee tee ee. 3
3. Conidiophores with a distinctive light-dark-light pigmentation pattern; conidia
brown to pale brown, 9=12exC7 1OUM aac ws oe ey ee ee B. africana
3. Conidiophores concolorous; conidia yellow, 6.0-7.4 x 6.4-8.6um .... B. triangularis
Acknowledgments
The authors are grateful to Drs David Minter and James Scott for their critical review
of the manuscript and suggestions for revision. The authors also want to express their
gratitude to, Pacific Environmental Consulting, North Vancouver, British Columbia, and
SanAir Technologies Laboratory, Powhatan, Virginia, for providing specimens.
Literature cited
Carmichael JW. 1955. Lacto-fuchsin: a new medium for mounting fungi. Mycologia 47: 611.
Ellis MB. 1976. More Dematiaceous Hyphomycetes. Commonwealth Mycological Institute, Kew.
Holmgren, PK, Holmgren NH. 1998. [continuously updated]. Index Herbariorum: A global
directory of public herbaria and associated staff. New York Botanical Garden's Virtual
Herbarium. http://sweetgum.nybg.org/ih/
Keshava Prasad TS, Bhat DJ. 2002. Stellomyces kendrickii, a new hyphomycete from India.
Mycotaxon 84: 61-63.
Kiffer E. 1973. Contribution a étude de la microflore fongique de Congo. II. Torula ndjilensis sp.
nov. et Balanium africanum sp. nov. Le Naturaliste Canadien 100: 257-263.
Kirk PM. 1985. New or interesting microfungi. XIV. Dematiaceous hyphomycetes from Mt Kenya.
Mycotaxon 23: 305-352.
Morgan-Jones G, Sinclair RC, Eicker A. 1987. Notes on Hyphomycetes. LII. Stellomyces, a new
denticulate genus from the Transvaal, South Africa. Mycotaxon 28: 447-451.
Whitton SR, McKenzie EHC, Hyde KD. 2002. Microfungi on the Pandanaceae: a revision of the
hyphomycete genus Balaniopsis with two new species. Mycoscience 43: 67-72.
Volume 105, pp. 111-117 July-September 2008
Radiigera tropica (Geastraceae, Geastrales),
a new species from a tropical rain forest of Thailand
TAKAMICHI ORIHARA’, TAIGA KASUYA?,
SOUWALAK PHONGPAICHIT? & YAOWALUK DISSARA?
h2-berg-3f@red.megaegg.ne.jp
‘Department of Agrobiology, Graduate school of Agriculture, Tottori University
4-101 Koyama-cho-minami, Tottori 680-8553, Japan
tkasuya@sakura.cc.tsukuba.ac.jp
Laboratory of Plant Parasitic Mycology
Graduate Shcool of Life and Environmental Sciences, University of Tsukuba
Tsukuba, Ibaraki 305-8572, Japan
souwalak.p@psu.ac.th
*Department of Microbiology, Faculty of Science
Prince of Songkla University, Hat Yai, Songkhla 90112, Thailand
Abstract — A new species, Radiigera tropica, is described and illustrated from a tropical
rain forest of Thailand dominated by Dipterocarpus spp. ‘The species is morphologically
characterized by a felty to floccose peridial surface, a four-layered peridium and small,
globose basidiospores (3-4.6 um diam) with low, dense verrucae. Moreover, some
specimens have minute, granular peridioles. Additionally, it shares characteristics of
both previously recognized subgenera, Taylorae and Radiigera. Following a taxonomic
description, reconsideration of these subgenera and characterization of the species
among previously described Radiigera species are discussed.
Key words — Geastrum, sequestrate fungi, gasteromycetes, taxonomy, Phallomycetidae
Introduction
Members of the genus Radiigera Zeller (Geastraceae, Geastrales) produce
hypogeous to subepigeous basidiomata that strongly resemble immature
basidiomata of Geastrum spp. but are macroscopically distinguishable from the
latter in that the basidiomata either do not open in a stellate manner or remain
closed at maturity. So far, four species of Radiigera have been recognized from
North America and Europe (Zeller 1944, Guzman 1971, Kers 1976, Dominguez
de Toledo & Castellano 1996). The crucial morphological difference is that even
when immature, Radiigera lacks both a well-developed endoperidium and an
112... Orihara & al.
apical pore, which are universally present in Geastrum spp. (Dominguez de
Toledo & Castellano 1996). Molecular phylogenetic analysis of three Radiigera
species with a concatenated five-locus dataset of nuclear and mitochondrial
DNA strongly supports nesting of Radiigera spp. within the Geastrum spp. clade
and implies that Radiigera, as currently recognized, is polyphyletic (Hosaka
2005, Hosaka et al. 2006).
In this paper, we describe and illustrate a new hypogeous fungus collected
from a tropical rain forest dominated by Dipterocarpus spp. in Songkhla
Province, Southern Thailand. We provisionally assign the species to Radiigera,
a genus that hitherto has not been reported from Asia, based on macro- and
micromorphological characters.
Methods
The specimens examined are deposited in the mycological herbaria of National
Museum of Nature and Science, Tokyo, Japan (TNS), Prince of Songkla
University, Hat Yai, Thailand (PSU) and Natural History Museum and Institute,
Chiba, Japan (CBM).
Fresh basidiomata collected in the field were observed macroscopically and
then air-dried for later examination. Free-hand sections of the specimens were
mounted in water, 5% KOH or Lactophenol-Cotton blue for standard light
microscopy. Each micromorphological structure was measured in Lactophenol-
Cotton blue on glass slides. Measurement of spores was from mature specimens.
Spore dimensions were based on the measurement of 50 randomly selected
spores. Color names are in general terms of the authors.
The surface features of spores were observed by scanning electron microscopy
(SEM). For SEM, gleba was dusted onto a specimen holder attached with
double-sided adhesive tape and then coated with platinum-palladium with
an E-1030 Ion Sputter Coater (Hitachi, Tokyo, Japan). The treated gleba was
examined with an S-4200 SEM (Hitachi, Tokyo, Japan) operating at 20.0 kv.
- Taxonomy
Radiigera tropica Orihara & T. Kasuya, sp. nov. Ficures 1-10
MycoBank MB 511725
Basidiomata globosa vel depresso-globosa, ca. 2.2 cm in diam., superficie coacta vel
floccosa, spadicea vel brunneola. Peridium 4-stratosum; stratum myceliale ochraceum,
ex hyphis crassitunicatis compositum, quae exterioribus laxe intertextis 2.5-6 um latis
interioribus dense intertextis 1-3 um latis compositum, cum contextu intimo amorpho
30-65(140) um crasso; stratum fibrosum 20-350 um crassum, ex hyphis crassitunicatis,
hyalinis, intertextis, 2.5-6.3 um latis compositum; stratum pseudoparenchymatum 280-
500 um crassum, ochraceum vel brunneum; endoperidium nullum vel depauperatum.
Gleba atro-brunnea, pulverulenta, saepe cum peridiolis subglobosis usque ad 150 um diam.
Capillitium 2.5-7 um latum, crassitunicatum, fuscum, non sinuosum, non septatum, cum
Radiigera tropica sp. nov. (Thailand) ... 113
granulis reliquis haerens. Basidiosporae globosae, verrucosae, 3-4.6 um diam., verrucis
minutis, densis.
Ho.Lotypus: THAILAND, Songkhla Province, Ton Nga Chang Waterfall, fere hypogaea
in silvis tropicali, 21 Feb.2005, leg. T: Kasuya (TNS-F-17629).
ETYMOLOGY: Latin, tropica refers to the characteristic distribution of the species.
BAsIDIOMATA hypogeous, globose to depressed globose, ca. 2.2 cm in diam.,
surface felty to floccose with debris, pale tawny to pale brown, thick hyphal
mat not observed in soil, basal rhizomorphs of pale brown, thick-walled (0.4-
0.7 um thick), filamentous hyphae parallel to surface with scattered, clamped,
non-ampullate septa, 1-2 um broad. PERip1uM four layered; exoperidium (=
mycelial layer) 200-450 um thick, tawny to pale brown, further divided into
three layers: the outer hyphal layer 80-220 um thick, acyanophilous, of thick-
walled (0.8-2.0 um thick), yellowish brown, aseptate, loosely interwoven
or partly perpendicularly arranged, filamentous hyphae, 2.5-6 um broad,
grading to inner; the inner hyphal layer 80-200 um thick, acyanophilous, of
thick-walled (0.4-1.4 tum thick), brown, aseptate, densely interwoven, narrow,
filamentous hyphae, 1-3 um broad, grading to inner amorphous stratum; the
innermost exoperidial layer (= amorphous stratum) 30-65(140) um thick,
cyanophilous, where the mycelia collapse in mature specimens; fibrous layer
20-350 um thick, acyanophilous, easily separable from the exoperidium, of
thick-walled (1-3 um thick), white, refractive, aseptate, loosely interwoven,
filamentous hyphae, 2.5-6.3 um broad; inner layer (= mesoperidium) 280-500
uum thick, of persistent, ochraceous to brown, thick-walled (0.7-2.6 um thick)
pseudoparenchymatous cells up to 25 um in diam.; endoperidium absent or if
present, poorly developed, comprised of pale grayish brown to reddish brown,
thick-walled (0.8-4 um thick), aseptate, densely interwoven hyphae, 3-10 um
broad. CoOLUMELLA indistinct, more or less columnar, white, collapse at maturity,
of thick-walled (0.5-0.8 um thick), branched, interwoven hyphae 1-5.5 um
broad, clamp-connections absent. GLEBA white at immaturity, then becoming
blackish brown, powdery at maturity, in some immature specimens comprised
of radially arranged cylindrical elements, ca. 1 x 0.1-0.2 mm, along with a mass
of granular peridioles, in other immature specimens never comprised of such
cylindrical elements or peridioles; cylindrical elements consisting of bundles of
tubular peridioles comprised of pale brown, palisade basidiole-like cells, and
tubular elements comprised of helicoid filamentous hyphae, 0.8-2 um broad
which partly loosen; The granular peridioles subglobose to depressed-globose,
15-30 x 20-52 um at immaturity, 40-150 x 25-100 um at maturity exposing
a blackish mass of basidiospores, comprised of hyaline to pale brown, palisade
cells deliquescing at maturity. CAPILLITIAL THREADS 2.5—7 um broad, more or
less bundled, thick-walled (0.8-2.8 um thick), pigmented with pale brown to
dark brown, non-branched, straight, non-septate, surface often covered with
114... Orihara & al.
adherent debris, appearing almost warty. BASIDIA not observed in peridioles;
in specimens not containing peridioles, sacciform to lecythiform, hyaline, 4-8-
spored, ca. 12-25 x 7-14 um, collapsing at maturity; generative hypahe hyaline,
thin-walled, filamentous, more or less sinuate, 0.8-5 um broad, with scattered
clamp connections, collapsing at maturity. BasipiosporEs globose, minutely
and densely warty, dark brown to blackish brown, 3-4.6 um in diam. (4 um in
mean diam.) including ornaments, basal apiculus not prominent, almost flat,
not forming ridges with the surrounding verrucae.
HaBiTaTt — Hypogeous, gregarious under Dipterocarpus spp.
DISTRIBUTION — Known only from Thailand (Songkhla Province).
SPECIMENS EXAMINED — THAILAND. SONGKHLA PROVINCE: Ton Nga Chang
Waterfall, February 21, 2005, T. Kasuya, Orihara 605, TNS-F-17629 (HOLOTYPE),
PSU-FB-21205 (IsoTYPE), CBM-FB-37171 (IsoTYPE); December 6, 2005, T. Kasuya, S.
Phogpaichit and Y. Dissara, Orihara 662, TNS-F-17661.
Discussion
Ecologically, Radiigera tropica occurs in tropical rain forest dominated
by Dipterocarpus spp. Although Dipterocarpus spp. are known as major
ectomycorrhizal host trees in the habitat, ectomycorrhizal association of
R. tropica has not yet been confirmed. Radiigera tropica is morphologically
characterized by the felty to floccose peridial surface with debris, the four-
layered peridium and the small basidiospores with low, dense verrucae.
Another notable feature is the presence of two glebal types: some specimens
have the radially arranged cylindrical elements and tubular or granular
peridioles within the gleba (Figs. 5-7), but others, as well as the remaining
Radiigera species, never contain these tissues (Fig. 2). The immature peridioles
of the former glebal type are comprised of palisade cells although we did not
observe any basidiospores within them in early developmental stages. It is
known that Schenella T. Macbr. (= Pyrenogaster Malencon & Riousset), a genus
recently transferred from Myxomycetes to Basidiomycetes (Estrada-Torres et al.
2005) and closely related to Radiigera, forms peridioles. However, the generic
characters of Schenella — such as smooth capillitium, filiform basidia and
absence of the fibrous layer in the peridium — were not observed in R. tropica.
Additionally, the mass of minute, granular, subglobose peridioles of R. tropica
Figs. 1-10. Radiigera tropica (from holotype). 1. Basidiomata. 2. Cross-section of a dried immature
basidioma without peridioles. 3. Peridium: A. Exoperidium. B. Fibrous layer. C. Inner layer -
(mesoperidium). D. Endoperidium. 4. SEM photomicrograph of a basidiospore with an indistinct
basal apiculus. 5. Glebal cylindrical elements along with minute, immature granular peridioles. 6.
Immature, granular peridioles. 7. Cylindrical elements in the immature gleba. 8. Basidiospores. 9.
Basidia. 10. Capillitium. Bars: Figs. 8-10 = 10 um.
Radiigera tropica sp. nov. (Thailand) ... 115
116 ... Orihara & al.
has little morphological affinity with the radially arranged, conical to pyramidal
peridioles of Pyrenogaster species. Therefore, we consider that there is not a
close relationship between R. tropica and Schenella spp. and that the placement
of R. tropica within the genus Radiigera is justifiable. Further molecular studies
may reveal a different phylogenetic relationship of the species.
Of all known taxa of, Radiigera, R. taylorii (Lloyd) Zeller most closely
resembles R. tropica in overall peridial structure and small (3-4 um
diam) basidiospore size but is distinguished by a smooth peridial surface,
perpendicularly arranged hyphae of an outermost exoperidial layer and less
densely arranged verrucae of basidiospores. Radiigera fuscogleba Zeller differs
from R. tropica in producing thin-walled exoperidial hyphae with clamp
connections and larger basidiospores (4.5-7(8) um diam, Dominguez &
Castellano 1996). The felty to cottony surface of R. bushnellii L.S. Dominguez
& Castellano reminds us of R. tropica, but R. bushnellii differs in having an
undifferentiated exoperidial structure, a poorly developed (sometimes absent)
capillitium and a more or less protruding basal basidiospore apiculus that forms
ridges with the surrounding verrucae. Radiigera flexuosa L.S. Dominguez &
Castellano is unique in having irregularly dehiscent basidiomata, a three-
layered peridium and thin-walled, contorted, sinuous capillitial threads, all of
which are not observed in R. tropica.
Dominguez de Toledo & Castellano (1996) recognized subgenus Taylorae for
R. taylorii, which was defined by small (3-4 um diam.), globose basidiospores,
a peridial mycelial layer of thick-walled hyphae perpendicularly arranged
towards the outside, and a peridial surface that is smooth and without adherent
debris. The other three species (ie., R. fuscogleba, R. bushnellii and R. flexuosa)
were placed in subgenus Radiigera Zeller. Radiigera tropica, however, shares
subgeneric characters of both subgenera Taylorae and Radiigera: relatively
small basidiospores and thick-walled exoperidial hyphae are characteristic
of subgenus Taylorae, whereas the felty to floccose peridial surface with
adherent debris and the loosely interwoven or less perpendicularly arranged
outermost exoperidial hyphae are characteristic of subgenus Radiigera.
Moreover, molecular phylogenetic analyses by Hosaka et al. (2006) indicate a
well-supported phylogenetic affinity between R. taylorii and R. bushnellii (i.e.
>0.95 Bayesian posterior probability; >70% maximum parsimony bootstrap
value), suggesting that the taxonomic boundary between subgenera Taylorae
and Radiigera is artificial. The phylogenetic study further suggests that the
genus Radiigera as currently delimited is artificial and that all species except-
the type (R. fuscogleba) should probably be assigned to Geastrum (Hosaka et
al. 2006, Hosaka pers. comm. 2008). Therefore, our assignment of R. tropica
to Radiigera is provisional, subject to taxonomic re-evaluation pending future
molecular analyses. In summary, reconsideration of both infra- and intrageneric
Radiigera tropica sp. nov. (Thailand) ... 117
classification in Radiigera with more detailed molecular methods would be
useful in the future.
Radiigera tropica is the first Radiigera species recorded from Asia and tropical
regions. Its presence suggests that Radiigera has wide distribution from tropical
to boreal areas. Although Radiigera is a small genus, more intensive studies in
Asia and its adjacent areas may reveal additional species diversity.
Acknowledgments
We are grateful to Dr. Ken Katumoto for checking the Latin diagnosis and giving us
helpful suggestions. We also acknowledge Prof. Nitaro Maekawa for critically reading
of the manuscript and providing valuable comments. We express our gratitude to Dr.
Michael Castellano and Dr. Kentaro Hosaka for reviewing the manuscript.
Literature cited
Dominguez de Toledo LS, Castellano MA. 1996. A revision of the genera Radiigera and Pyrenogaster.
Mycologia 88: 863-884.
Estrada-Torres A, Gaither TW, Miller DL, Lado C, Keller HW. 2005. The myxomycete genus
Schenella: morphological and DNA sequence evidence for synonymy with the gasteromycete
genus Pyrenogaster. Mycologia 97: 139-149.
Guzman G. 1971. Notas sobre los géneros Radiigera and Mesophelliopsis en México. Bol Soc Méx
Micol 5: 7-11.
Hosaka K. 2005. Systematics, phylogeny, and biogeography of the Hysterangiales and related taxa
(Phallomycetidae, Homobasidiomycetes). Ph.D. Thesis, Oregon State University, Corvallis,
Oregon. 361 pp.
Hosaka K, Bates ST, Beever RE, Castellano MA, Colgan III W, Dominguez LS, Nouhra ER, Geml J,
Giachini AJ, Kenney SR, Simpson NB, Spatafora JW, Trappe JM. 2006. Molecular phylogenetics
of the gomphoid-phalloid fungi with an establishment of the new subclass Phallomycetidae and
two new orders. Mycologia 98: 949-959.
Kars L. 1976. Radiigera Zeller, a genus of Gasteromycetes new to Europe. Botaniska Notiser 129:
173-178.
Zeller SM. 1944. Representatives of the Mesophelliaceae in North America. Mycologia 36:
627-637.
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MYCOTAXON
Volume 105, pp. 119-136 July-September 2008
A new genus, Cruentomycena,
with Mycena viscidocruenta as type species
RONALD H. PETERSEN’, KAREN W. HUGHES’, EDGAR B. LICKEY?,
ALEXANDER E. KOVALENKO?, OLGA V. MOROZOVA3
& NADEZHDA V. PSURTSEVA3
' repete@utk.edu
' University of Tennessee, Knoxville, TN 37996 USA
’ Bridgewater College, Bridgewater, VA, 22812 USA
> Komarov Botanical Institute, St. Petersburg, 197376, Russia
Abstract — Mycena viscidocruenta is presently known only from the Antipodes.
Collections from far eastern Russia were shown to be closely related both by morphology
and by sequence homology. Phylogenetic analyses based on ribosomal LSU and ITS
sequences show that the M. viscidocruenta clade is sister to Panellus stypticus and
Panellus (Dictyopanus) pusillus, two laterally stipitate taxa, in the panelloid clade.
Resinomycena rhododendri, the type species of Resinomycena is basal to this group.
A new genus, Cruentomycena, is proposed to accommodate C. viscidocruenta (typus
generis) and a new species, C. kedrovayae.
Key words — biodiversity, biogeography
Introduction
Mycena viscidocruenta is a small (cap 7-11 mm), bright red agaric with a
very slimy stem and cap. Fruiting on leaves and small woody debris, it has
been reported from the southeastern Australian mainland and Tasmania
(Grgurinovic 2002; Fungimap, Australia: http://fungimap.rbg.vic.gov.au), and
from the North and South Islands of New Zealand (McKenzie et al. 2000, on
Nothofagus debris; Segedin & Pennycook 2001; E. Horak, pers. com.). An LSU
rDNA sequence of Tasmanian material was used by Moncalvo et al. (2002), who
showed that collection to belong to the panelloid clade (denoted as /panelloid).
That paper further noted that a relationship among Resinomycena rhododendri
(typus generis), Mycena viscidocruenta, Panellus (pleurotoid, lamellate) and
Dictyopanus (pleurotoid, poroid) had not previously been reported (Singer
1986), but did not discuss morphological circumscription of M. viscidocruenta
or the proposed merger of Dictyopanus and Panellus (Burdsall & Miller 1975).
120 ... Petersen & al.
Two collections of a similar agaric from the Primorsky Territory in far eastern
Russia on Alnus catkins were thought to greatly extend the geographic range
of the species (Petersen & Hughes 2007); the known distribution of M.
viscidocruenta does not include temperate east Asia or any geographical region
between Russia and southern Australia/New Zealand.
The identity of the source of an LSU DNA sequence (Moncalvo et al. 2002)
was insecure, and the temperate Russian collections, while molecularly close
to Tasmanian material, seemed undescribed. This paper takes up this small
problem with the following intent: 1) redescription of Mycena viscidocruenta
from additional Australian specimens including the type; 2) description of the
Russian taxon; 3) discussion of the molecular placement of these collections;
and 4) proposal of a new genus name to accommodate these specimens.
Methods and materials
Morphological analyses
In order to associate morphological features with DNA sequences, specimens from
Kew (K) and Adelaide (AD) were borrowed and examined (see below). All observations
were with Phase Contrast optics (PhC), with magnification up to 1200x on an Olympus
BX60 research microscope. Colors within quotation marks are from Ridgway (1912);
those cited alphanumerically are from Kornerup & Wanscher (1967).
Molecular protocols
For DNA extractions from herbarium specimens, a small piece of gill tissue was
removed and ground to a fine powder with sterile grinding sand. The mixture was added
to a CTAB extraction buffer (Carlson et al. 1991) and extracted as described in Mata et
al. (2007) or extracted using a Qiagen DNA easy Plant Mini Kit following manufacturer's
instructions. PCR amplification of the ribosomal ITS1-5.8S-ITS2 region was carried out
with primers ITS1F and ITS4. PCR of the ribosomal Large Subunit 3’ end was carried
out with primers LR7 (Moncalvo et al. 2000) and ITS3. DNA sequencing followed
manufacturer's directions for Big Dye Terminator mix (ABI) with forward primer ITS5
and reverse primer ITS4 for the ribosomal ITS region and primers BR5, LR21 and LROR
for the LSU region. Primers ITS3, ITS4 and ITS5 are described in White et al. (1990);
Primer ITS1F is described in Gardes & Bruns (1993); Primers LR5, LR21 and LROR are
given in Moncalvo et al. (2000). Sequences were aligned and compared using the GCG
suite of programs (GCG 2000). Sequences obtained for this study are given in Table 1.
Molecular Data Analyses
Initial sequence alignments were carried out using the “pileup” program in GCG
which performs UPGMA, followed by manual adjustments to the alignment. For both
ITS and LSU regions, Mycena (Prunulus) pura was selected as an outgroup based on
Moncalvo et al. (2002), on blast homology in Gen Bank and on preliminary parsimony
analyses which included other Mycena and Favolaschia species. There were extensive
but informative gaps in ITS sequences of Panellus stypticus collections from Australia
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a ke a a. every BIIOISIA “elpeajsny fT ‘ds (snupdodgaiq) snyjating 6690. Sa = Sen i x9CPIOTAV
Ties see ae ee A ca
Se ree SATIANVY
‘papnjouod ‘J ajquy
Cruentomycena gen. nov. for Mycena viscidocruenta ... 123
and New Zealand (Jin et al. 2001b). For this ITS data set, gaps were treated as missing
in maximum parsimony analysis. An additional 172 bases representing four regions of
the ITS area were deleted in all analyses because alignment of outgroups with ingroups
was uncertain. Gaps were minimal but informative in the LSU data set and maximum
parsimony was performed with gaps treated as a fifth base.
Maximum parsimony was performed using PAUP* (Swofford 2001). Characters
were unordered and weighted equally. Trees were limited to 100; multistate taxa were
interpreted as uncertainty; starting trees were obtained via stepwise addition; addition
sequence = furthest; the number of trees held at each step during stepwise addition =
1; branch-swapping algorithm was tree-bisection-reconnection (TBR); steepest descent
option was not in effect; maxtrees setting = 100. One-thousand bootstrap replicates
were performed. Modeltest was used to determine the appropriate model of evolution
for Bayesian analysis (Posada & Crandall 1998). The model selected by Model Test
was a general time reversible model with rate variation (gamma) and both variable
and invariant sites (GTR+G+I). Bayesian analysis was performed using Mr Bayes
(Huelsenbeck & Ronquist 2000) with the following settings; nst = 6, basefreq = estimate.
The MCMC search was run with 4 chains for 500,000 generations with sampling every
100 generations. The first 1,000 trees were discarded and “burnin” based on preliminary
analyses showing that likelihood values had reached stability with the first 1000 trees.
Posterior probabilities were estimated by sampling trees generated after likelihood
values diverged.
Results
Taxonomic description
Cruentomycena R.H. Petersen, Kovalenko & O. Morozova, nom. et stat. nov.
MyYcoBANK MB 511635
Bastonym: Mycena sect. Viscidocruentae Maas Geest. & Hauskn. 1998. Osterr. Zeitschr.
Piz FAD,
TYPE SPECIES: Cruentomycena viscidocruenta (Cleland) R.H. Petersen & Kovalenko.
ETYMOLOGY: cruentus = splashed with blood; Mycena = resembling Mycena.
Basidiomata mycenoidea vel marasmielloidea. Pileus diminutivus, planus ad depressus,
viscidus, sanguineus; trama pseudoparenchyma, pseudoamyloidea. Lamellae arcuatae ad
decurrentae, crassae, cinnabar-sanguineae, marginatae. Stipes tenuis, levis, aequalibus.
Hab: in ramunculi et folia. Hyphae pileipellis angustae, tenuitunicatae, fibulatae.
Cheilocystidia praecentia. Pleurocystidia nulla. Sporae elongatae, amyloideae.
Discussion: In proposing a new genus name, elevation of the section name
would have produced a tautonym, Viscidocruenta viscidocruenta, illegitimate
under the ICBN [(Vienna) Art. 23.4]. A new genus name is therefore
necessary.
Cruentomycena viscidocruenta (Cleland) R.H. Petersen & Kovalenko, comb. nov.
MycoBANK MB 511636
Basionym: Mycena viscido-cruenta Cleland. 1924. Trans. Proc. Roy. Soc. S. Australia
48: 239. [nom. nov. for Mycena coccinea Cleland & Cheel. 1919. Trans. Proc. Roy.
124 ... Petersen & al.
Soc. S. Australia 43: 284, nom. illeg., non M. coccinea (Sowerby) Quel. 1880 [1879].
Bull. Soc. Amis Sci. Nat. Rouen II 15: 155].
LECTOTYPE: (design. Grgurinovic 1997): AUSTRALIA, New South Wales, Mosman, S
33° 14, E 150° 50’ (precn. 3), 24.V.1914, coll. J.B. Cleland (as Mycena coccinea), det.
C.A. Grgurinovic (as M. viscidocruenta), (JBC) s.n. (AD-C12182).
EXEMPLAR — AUSTRALIA, Victoria, Sherbrooke Forest, Hardy Creek, S$ 37° 53°43”,
E 145° 21’ 44” (precn 3), 2.1V.1981, coll. & det. CA Grgurinovic, CAG 9481 (AD-
C12334).
COLOR ILLUSTRATIONS — Cleland & Cheel (1919), Grgurinovic (1997). http://www.
landcareresearch.co.nz.
MACROCHARACTERS — BASIDIOMATA (Fig. la) small, mycenoid; PILEus 4-
11(-14) mm broad, plane to shallowly convex, centrally depressed or almost
umbilicate, viscid to glutinous when fresh, laccate to matt when dried, rich
reddish crimson, blood red, minutely rugulose, outward gently crenate, deeply
striate, color red-black upon drying; margin subcrenate when fresh, sometimes
appearing striate and often inrolled when dried; LAMELLAE subdistant, adnate
with decurrent tooth to arcuate, terminating quite abruptly on stipe, thick,
slightly paler than cap (rose color of Cleland) and becoming dark cinnabar to
pallid gray-mauve on drying, very minutely denticulate, lamellar edges very
slightly darker than lamellar face when fresh, distinctly marginate to blood red
or deep purple upon drying; StrPE 6-25 mm long, 0.4-1 mm broad, terete,
equal, glutinous to viscid, blood red, hollow, perhaps producing minimal
blood-red juice upon bruising; base sometimes swollen into a hemisphere
or a rounded carbuncle concolorous with stipe, occasionally with superficial
cinnabar tomentum. Red pigment leaching copiously in formalin (Cleland)
and KOH.
Hasitat — On fallen twigs and associated leaf fragments; “deep shady places”
(Cleland).
MICROCHARACTERS — PILEIPELLIS with superficial viscid layer (with
adherent spores) covering radially oriented, thin-walled (easily overlooked),
unornamented hyphae 4-9 «tm diam., hardly inflated, not encrusted,
frequently septate and occasionally demonstrably clamped; terminal cells
undifferentiated, simply rounded apically. Pileus trama of significantly inflated
hyphae up to 28 um diam., thin-walled, constricted at septa, clamped, almost
pseudoparenchymatous, perhaps weakly pseudoamyloid; Basip1a (Fig. 1b)
20-25 x 6-8 um, clavate, clamped, thin-walled, 4-spored; sterigmata persistent
after spore discharge, awl-shaped, stiff; BAstp1ospores (Fig. 1c) amyloid,
(6.5-)8-10 x (2.5-)3-4.5 um, thin-walled, elongate pip-shaped, tapering
asymmetrically toward hilar appendage, aguttulate, hyaline. Lamellar trama
irregular, of somewhat inflated cells, weakly pseudoamyloid (but difficult
to distinguish from merely pink-pigmented; CHriLocystipia (Fig. 1d,e)
scattered along lamellar edge, dimorphic: 1) (Fig. 1d) 17-42 x 11-24 um,
Cruentomycena gen. nov. for Mycena viscidocruenta ... 125
Fig. 1. Cruentomycena viscidocruenta. AD-C12334.
a. Basidiome, x6. b. Basidiospores. c. Basidia. d, e. Cheilocystidia. f. Caulocystidia.
Standard bar = 10 um for basidiospores and basidia, otherwise = 20 um.
sphaeropedunculate to almost sessile, very thin-walled; and 2) (Fig. le) 27-41
x 7-14 um (at widest point), ten pin-shaped or sublecythiform with narrowed
neck and slightly expanded capitulum (6-7 um broad). Stipe surface covered
with thin viscid layer; hyphae strictly parallel, thin-walled, 3.5-7 um diam.,
infrequently septate, ?clamped; occasionally producing out-turned hyphal tips;
CAULOCYSTIDIA (Fig. 1f) somewhat inflated, 28->75 x 8-14 um, thin-walled,
of undetermined origin; contents homogeneous, with very pale pinkish tint.
126 ... Petersen & al.
COMMENTARY — At least three of the collections listed below were in the
hands of Cleland at the time he and Cheel proposed M. coccinea, and therefore,
together with additional specimens at AD not examined, constitute a group of
syntypes, from which Grgurinovic (1997: 261) chose a lectotype (see above).
Fungus material remaining in the lectotype, however, is meager, so a more
recent collection (AD-C12334) has been cited as “exemplar.” The designation
“epitype” is not used here, in hopes that a future, more appropriate collection
can be employed to extract DNA for comparison.
We have seen no specimen comprising more than three basidiomata,
implying that when found in nature, only a few basidiomata are produced by a
single, localized mycelium.
Macromorphologically, the specimens of C. viscidocruenta (including those
labeled as M. coccinea) are nearly inseparable from the Russian material: colors
of pileus, lamellae, lamellar edge, and stipe all match. Size ranges of basidiomata
from the two regions were also similar, as were basidiospores, basidia and
caulocystidia. although habitat clearly differs. Cleland & Cheel (1919) noted
that color leached from tissues in formalin, and the phenomenon is also strong
in KOH. Conversely, Russian material exhibits only one type of cheilocystidia
while Australian specimens clearly produce two types mixed over the lamellar
edge, and cheilocystidia of similar shape are smaller in the Russian material.
Spores are comparable, although rough measurements might indicate that in
Russian specimens they are slightly wider. Russian material was not obviously
viscid when collected even though it had been raining for several days, but
Maas Geesteranus & Hausknecht (1998) briefly discussed discrepant reports on
this character, including that by Grgurinovic & Holland (1982).
Grgurinovic (1997) placed M. viscidocruenta in sect. Hygrocyboideae (Fr.)
Singer, with further placement in subsect. Gummosae (J.E. Lange) Oort.
Maas Geesteranus & Hausknecht (1998) recognized the species as unique,
and proposed sect. Viscidocruentae to accommodate it. Grgurinovic (2002)
examined numerous specimens, citing its distribution as south-eastern
Australia, Queensland and New Zealand. Neither Grgurinovic (1997, 2002)
nor Maas Geesteranus & Hausknecht (1998) anticipated that DNA sequences
would show it to be even less related to the bulk of Mycena, and a candidate for
genus rank.
Cleland made reference to a specimen in formalin (note with AD-C12187)
but this specimen has been lost. The same specimen was said to resemble
“Miss Clarke Fig., A3 [believed to be on p. 4 of Cleland folder]” a reference to
a watercolor rendering thought to still exist in a folder of Cleland illustrations
at AD. This watercolor may have been published as Cleland & Cheel (1919: Pl.
29, Fig. 4).
Cleland furnished spore measurements, which closely match those reported
above. (Cleland: 7-8.2 x 2.5-3.4 um; 8.5 x 3.5 um; 8 x 3.8 um).
Cruentomycena gen. nov. for Mycena viscidocruenta ... 127
SPECIMENS EXAMINED — AUSTRALIA, New SoutH WALES, Mosman, § 33° 14,
E 150° 50} 21.VI.1914, coll. JB Cleland (as M. coccinea), s.n. det. C.A. Grgurinovic
(as M. viscidocruenta). AD-C12187 (formerly ADW 12195, “type”, later labeled as
“syntype’); Mosman, S 33° 14} E 150° 50’ (precn. 3), 24.V.1914, coll. J.B. Cleland (as
Mycena coccinea), det. C.A. Grgurinovic (as M. viscidocruenta), (JBC) s.n. (AD-C12182;
lectotype); SouTH AUSTRALIA, Southern Lofty, Waterfall Gully, S 34° 57’ 43”, E 138° 40°
17” (precn: 3), 19.VI.1921, coll. J.B. Cleland (as M. coccinea), det. C.A. Grgurinovic (as
M. viscidocruenta), ADW 12201 (AD-C12318); TASMANIA, Mount Wellington, Myrtle
Gully, 25.111.2003, coll. G. Gates & D.A. Ratkowsky G&R 0368, K(M) 117458); VICTORIA,
Sherbrooke Forest, Hardy Creek, S 37° 53°43”, E 145° 21 44” (precn 3), 2.IV.1981, coll.
& det. CA Grgurinovic, CAG 9481 (AD-C12334); VicrortA, Sherbrooke, Hardy Creek,
S 37° 53° 43”, E 145° 21’ 44” (precn 3), 30.11.1981, coll. & det. C.A. Grgurinovic, CAG
11381 (AD-C12256); Vicrorta, Glen Nayook, S 37° 55’ 22”, E 145° 56’ 43” (precn 3),
15.IV.1981, coll. & det. C.A. Grgurinovic, CAG 20481 (AD-C12277).
Cruentomycena kedrovayae R.H. Petersen, Kovalenko & O. Morozova, sp. nov.
MycoBANK MB 511637 FIG. 2
Ut C. viscidocruenta, vel stipite et pilei vix glutinosa, habitat in Alnus amenta et
ramunculi, cheilocystidia uniformiter lato-clavata, et distributo geographica in Asia
occidentalis-septentrionali.
HoLotype — RUSSIA, PrRimorsky TERRITORY, Khasansky Dist., Kedrovaya Pad
Nature Reserve, N 43° 05.8693 E 131° 33.572) 104 m elev., 18.VIII.2005, coll. RHP & A
Kovalenko (as Xeromphalina sp.), TFB 11844 (TENN 60729).
EtyMoLocy — Denotes Kedrovaya Pad Reserve, Khasansky Dist., Primorsky Territory,
Russia, type location.
MACROCHARACTERS — BASIDIOMATA (Fig. 2a,b) small, marasmielloid;
PitEus 5-9 mm broad, shallowly convex, plane with central depression, to
subinfundibuliform, outward subtly crenate-striate, appearing glabrous but
unglazed, inward “Madder brown” (10E8), outward “Pompeian red” (10C6),
leaching copious carmine pigment into KOH; margin sometimes subcrenate,
deeply striate; LAMELLAE arcuate to decurrent, thick (almost fold-like),
occasionally forked, near “dragon's blood red” (8C8) to deep maroon, shallow;
lamellar edge minutely marginate (especially when dried: faces grayish maroon,
margin minutely deep, clear maroon); STIPE 8-13 x 0.5-0.8 mm, terete, slender,
tough, near “madder brown” (10E8) here and there glazed, perhaps as though
with a thin viscid layer, profoundly hollow; basal pad deep cinnabar red, not
widely spread, felty. 3% KOH on dried pileus turns instantly red-black; ODOR
none; TASTE none.
HaBitat — On Alnus catkins, Alnus twigs, and closely associated dead grass
stems.
MICROCHARACTERS — PILEIPELLIS with superficial, thin viscid layer; tissue of
repent, radially arranged hyphae 3.5-6.5 um diam., occasionally septate (and
occasionally obscurely clamped), unornamented, thin- to firm-walled (wall
never more than 0.5 um thick), apparently hyaline. Pileus trama dextrinoid,
128 ... Petersen & al.
constructed of intricately interwoven, thin- to firm-walled (wall never more
than 0.5 um thick), occasionally inflated (up to 30 um diam.), obscurely
clamped hyphae; slender (2.5-4 um diam.), frequently branched, thin-walled,
clamped hyphae occasional; vascular hyphae occasional, 4-6 um diam., thin-
walled, refringent (PhC). Lamellar trama interwoven, dextrinoid; tramal
hyphae uniformly 6-7 um diam., often asymmetrically inflated to 15 um diam.,
thin-walled, conspicuously clamped; vascular hyphae rare, 3.5-5 um diam.,
refringent (PhC); Basrp1A (Fig. 3a) 18-25 x 6-7 um, digitate to narrowly
clavate, clamped, 4-spored; contents with a few small, scattered guttules.
Basidioles digitate, not mucronate, prominently clamped; BAsip1osPporEs (Fig.
3b) 6-8(-9) x 3.5—4 um, ellipsoid, somewhat tapered proximally abaxially (i.e.
asymmetirically lacrymoid; narrowly pip-shaped), thin-walled, hyaline, weakly
amyloid; contents with 1-few, small, scattered guttules. Lamellar edge virtually
sterile, a solid beard of cheilocystidia; CHEILOcysTIpDIA (Fig. 3c) 21-59 x 7-14
um, pedicellate, broadly clavate, clamped, hyaline, thin- to firm-walled (wall
never more than 0.5 um thick), smooth (without ornamentation) or rarely with
some asymmetric outgrowth. Stipe surface naked, very slightly viscid; surface
hyphae 2-6 um diam., thin-walled, conspicuously clamped, repent, probably
adherent to underlying tissue; outer stipe flesh hyphae 6-11 tm diam., short-
celled (cells 20-50 um long), thin-walled, not constricted at septa, obscurely
clamped, dextrinoid; surface hyphae rarely produce out-turned terminal cells;
CauLocysTipiA (Fig. 3d), 32-75 x 5-17 um, very thin-walled, broadly clavate,
probably clamped, hyaline or very pale pink.
CuLTURE — LE(BIN) 2000. On malt extract agar mycelium covering 90
mm in 3.5 weeks. Advancing zone appressed; outline even. Mat wooly with
thin, radiating hyphae, matted, mealy white with age; odor absent; reverse
unchanging. Vegetative hyphae (2—-)2.5-5(-5.5) um diam. with frequent
prominent clamp connections. In age, colony kept under light may form
mycelial crust; interlocking hyphae form numerous swellings and round
cells. Laccase reaction = ++. Culture LE(BIN) 2000 maintained in Komarov
Botanical Institute Basidiomycetes Culture Collection LE (BIN) as subculture
on ale-wort agar slants and in vials under distilled water.
COMMENTARY — When collected, field characters did not satisfactorily place
basidiomata taxonomically. Almost all basidiomata were on Alnus catkins,
peduncles and twigs, with only a few on closely associated dead grass stems
and leaves. With decurrent, thick lamellae almost fold-like and small cinnabar _
basal pad, the specimens were noted as possibly Xeromphalina, although the
deep red colors were discordant for that genus. Filamentous black rhizomorphs
(now thought probably not associated with the basidiomata) further suggested
Xeromphalina
Cruentomycena gen. nov. for Mycena viscidocruenta ... 129
Fig. 2. Cruentomycena kedrovayae, basidiomata.
Holotype. a = x3. b = x7. Photos by A. Kovalenko.
130 ... Petersen & al.
Fig. 3. Cruentomycena kedrovayae, holotype.
a. Basidiospores. b. Basidia. c. Cheilocystidia, d. Caulocystidia.
Standard bar = 10 um for basidiospores and basidia; otherwise = 20 tum.
Later, ITS and LSU DNA sequences placed the collections together with
Panellus, Dictyopanus and Resinomycena in the panelloid clade (Moncalvo
et al. 2002). While internally macromorphologically anachronistic, the clade
was characterized by coralloid “dendrohyphidia,’ especially as cheilocystidia,
dextrinoid tramal tissues and amyloid spores. Basidiomata of the Russian °
collections failed to demonstrate the diagnostic oleocystidia and dendrohyphidia
of Resinomycena, so placement in this genus was abandoned.
Another taxon that Moncalvo et al. (2002) placed in this clade was Mycena
viscidocruenta. Basidiomata were diagnosed with bright red coloration, with
Cruentomycena gen. nov. for Mycena viscidocruenta ... 131
viscid pileus and stipe surfaces. When Australian material of M. viscidocruenta
was examined, microscopic characters similar to the Russian collections were
observed and ITS sequences virtually matched those of the Russian material
(2.86% base-pair difference). The collection used to generate the LSU sequence
for Moncalvo et al. (2002) was a Vilgalys collection from Tasmania, and LSU
sequences of the Russian collections matched that of the Vilgalys collection. We
considered M. viscidocruenta to be a likely identity of the Russian material, at
least congenerically.
In order to better understand taxonomic separation of the collections of M.
viscidocruenta from Resinomycena, we examined specimens of R. rhododendri
(typus generis) from herb. TENN. In all cases, abundant oleocystidia and
coralloid hyphal termini were observed on all surfaces of basidiomata (fully
described and illustrated by Redhead & Singer 1981; Redhead & Nagasawa
1987). Basidiomatal features were significantly different; slender, gracile, off-
white but clitocyboid in Resinomycena; shorter, hardly gracile, marasmielloid
and blood red in M. viscidocruenta. Also significant, however, was that two ITS
sequences representing R. rhododendri were basal to sequences representing
M. viscidocruenta collections at both ITS and LSU resolutions. At the LSU level,
the placement of the single sequence of R. acadiensis is unexplained and an ITS
sequence is unavailable for this taxon. Thus, morphologically and based on in-
hand material, M. viscidocruenta seemed discordant in clade /panelloid. A new
genus seemed necessary to accommodate M. viscidocruenta and the Russian
collections.
SPECIMENS EXAMINED (Resinomycena rhododendri) — USA, NorTH CAROLINA,
Haywood Co., GREAT SMOKY MOUNTAINS NATIONAL PARK, Cataloochee Cove, Caldwell
Creek Campground, 30.V1.1989, coll. DE Desjardin, TFB 1852 (TENN 48598); Swain
Co., GREAT SMOKY MOunrTaAINS NATIONAL Park, Mingus Mill Creek Trail, 13.VII.2006,
coll. E Lickey (as Clitocybe sp), TFB 13275 (TENN 61536). TENNESSEE, Blount Co.,
GREAT SMOKY MouNTAINS NATIONAL PARK, Elkmont, Jakes Creek Trail, 3.VII.1991,
coll. SA Gordon, RHP, DE Desjardin, V Antonin, TFB 3574 (TENN 50793); Knox Co.,
KNOXVILLE, Timberlake Rd., 3. VIII.1958, coll. L.R. Hesler, (TENN 23102).
SPECIMENS EXAMINED (C. kedrovayae) — RUSSIA, PRIMORSKY TERRITORY,
Khasansky Dist., Kedrovaya Pad Nature Reserve, N 43° 05.869, E 131° 33.572; 104 m
elev, 18.VIII.2005, coll. RHP & A Kovalenko (as Xeromphalina sp.), TFB 11844 (TENN
60729; holotype); same location, N 43°05.875, E 131°33.186, 18.VIII.2005, LE212084
[culture: LE(BIN)2000]; same location, N 43° 05.8697, E 131° 33.5723 16.VIII.2005, coll.
A. Kovalenko, TFB 11831 (TENN 60716).
MOLECULAR ANALYSES — ‘The results of Parsimony and Bayesian analyses
of ITS and LSU data sets were consistent (Figs. 4, 5). Both analyses place
Resinomycena rhododendri basal to Cruentomycena and to Panellus stypticus
and Panellus (Dictyopanus) pusillus. This relationship was also noted by
Moncalvo et al. (2002) who placed R. rhododendri basal to the panellioid clade
based on ribosomal LSU sequence data.
132 ... Petersen & al.
Discussion
Collection TENN 60729 (Cruentomycena kedrovayae) was largely
indistinguishable from collection G&R0368 [K(M): 117458] (Cruentomycena
viscidocruenta). The bright red hyphal contents easily dissolved in water and
KOH when tissue was rehydrated; spore shape was identical and spore size
nearly so. Cystidial shapes were identical and included some digitate forms;
pileus micromorphology was the same; pileocystidia were absent.
Resinomycena was proposed for species formerly in Mycena, Marasmius and
Omphalina that share digitate oleocystidia and dendrohyphidioid: elements
over pileal and stipe surfaces, and produce amyloid basidiospores and oleiferous
cheilocystidia (Redhead & Nagasawa 1987). Resinomycena rhododendri was
characterized as having oleiferous hyphae but hyphae lack the red contents of
M. viscidocruenta. Spores of R. rhododendri and C. viscidocruenta are similar in
shape and are hyaline, amyloid and smooth. Cheilocystidia are similar in shape
to those seen in C. viscidocruenta except that digitate cheilocystidia are seen in
C. viscidocruenta and were not reported for R. rhododendri. The habitat of R.
rhododendri and C. viscidocruenta is similar; leaf litter, twigs, and woody fruit
husks (TENN 60729 was fruiting on Alnus catkins, leaf litter and twigs).
Mycena viscidocruenta sensu lato is embedded in the panelloid clade
comprising Panellus stypticus and species traditionally within Dictyopanus (D.
pusillus and D. copelandii) (see also Matheny et al. 2006). Panellus pusillus was
transferred from Dictyopanus to Panellus by Burdsall & Miller (1975) based
on a number of shared morphological characters including lignicolous habitat,
ovoid (sometimes narrowly ovoid) amyloid spores, a basidiome with a well-
developed lateral stipe, a dry cuticle of densely entangled hyphae penetrated
by pileocystidia with refractive contents and bioluminescence. The proposed
relationship between P. stypticus and P. (Dictyopanus) pusillus was confirmed
using molecular data by Jin et al. (2001a). Panellus stypticus is widely distributed
in Northern and Southern Hemisphere temperate forests while P pusillus
is tropical or subtropical. Collections of P stypticus collected in Kedrovaya
Pad belonged to the Asian/US Pacific Northwest clade identified by Jin et al.
(2001b).
The finding of C. kedrovayae in temperate forests of eastern Russia produces
an apparent disjunct distribution (Petersen & Hughes 2007). Lack of reports of
this brightly colored species from China, Japan, southeastern Asia, Indonesia
and Malaysia may be due to its absence in these geographical regions (but see -
Corner 1994a: 239, Pl. 23, Fig. 28, for description of Mycena. roseilignicola
Corner). More likely, it has been collected under a different epithet, or is rare
and has been missed by collectors. A number of other taxa or Asian clades
of taxa share similar distributions including Lentinula edodes (Hibbett 1998),
Cruentomycena gen. nov. for Mycena viscidocruenta ... 133
AF042606 Mycena rutilanthiformis*
DQ457688 Mycena afn. pura Massachusetts Clone 1*
DQ457689 Mycena afn. pura Massachusetts Clone 2*
97/0 # AF261415 Resinomycena rhododendri USA, VA*
3674 Resinomycena rhododendri USA, TN, GSMNP
AF518634 Panellus stypticus*
8$3/0.90| AF261427 Paneilus stypticus*
AY014292 Panellus stypticus Switzerland
49/0.91 AY398745 Panellus stypticus Germany*
AF042638 Resinomycena acadiensis*
47/0.83
eae al
AF 261426 Panellus (Dictyopanus) sp.
AF261425 Panellus (Dictyopanus) pusillus
AY014291 Panelius (Dictyopanus) pusillus
11831 Cruentomycena kedrovayae Russia, sp. nov.
AF 261414 ‘Mycena viscidocruenta’ Australia, Tasmania*=
97/ Cruentomycena viscidocruenta gen. nov.
~~ 4 changes per site GR0368C3 Cruentomycena viscidocruenta gen. nov.
Australia, Tasmania
‘ Sequences not generated by the authors
Fig. 4. One of 100 trees derived from Parsimony analysis of sequences from the ribosomal ITS1-
5.8S-ITS2 region (maxtrees = 100 and was not increased). Bootstrap support for 1000 trees
followed by Bayesian posterior probabilities is given to the left of a node. Tree length was 182 steps.
All characters were unordered and equally weighted; 391 characters were constant. 33 variable
characters were parsimony-uninformative; 98 were parsimony-informative. Gaps were treated
as “missing.” Starting tree(s) obtained via stepwise addition. Addition sequence = furthest. Tree
length = 182; Consistency index = 0.8681, Homoplasy index = 0.1319.
Panellus stypticus (Jin et al. 2001b), Xerula globospora (Petersen & Nagasawa
2005) and Oudemansiella altissima (Corner 1994b).
134 ... Petersen & al.
DQ490643 Mycena afn. pura USA, Massachusetts*
EUS17506
EU517504
Mycena pura
Eastern Russia
EU517505
EU517507
400/1.00| EU517508 Resinomycena rhodedendri , USA
FU517509 Great Smoky Mountains National Park
EU517510
EU517515
EU517516 Cruentomycena viscidocruenta
94/1.00) EU517517 Australia, Tasmania
24/1.00 EU517518
EU517514
EU517511 Cruentomycena kedrovayae
£U517512 Russia, Primorsky Dist., Kedrovaya
EU517513
400/1.00) AY014291
L43376
143375
Panellus pusiilus
Southeastern USA
AF289062
AF289063
EU517519
AF 289064 Panellus stypticus
£U517520 Southern Hemisphere
£U517521
EU517522
EU517523
AF289070
AF289071
EU517528
EU517530 Panellus stypticus
EU517531 Eastern USA and Canada
EU517532
EU517533
EU517534
EU517529
AB084488
AF289065
AF 289066 Panellus stypticus
AF 289067 Asia/Western USA
EU517526
EU517527
AF289068
AY014292 Panellus stypticus
EU517524 Europe
EU517525
98/1.00
87/1.00
$6/1.00
_1 changes per site
*Not generated by authors
Fig. 5. One of 100 trees derived from Parsimony analysis of sequences from the ribosomal large
subunit gene (maxtrees = 100 and was not increased). Bootstrap support for 1000 trees followed by
Bayesian posterior probabilities is given to the left of a node. Tree length was 94 steps. All characters
were unordered and equally weighted; 816 characters were constant. 31 variable characters were
parsimony-uninformative; 45 were parsimony-informative. Gaps were treated as “fifth-base”
Starting tree(s) obtained via stepwise addition. Addition sequence = furthest. Tree length = 182;
Consistency index = 0.7619, Homoplasy index = 0.2381.
The percent ITS sequence difference between Kedrovaya Pad and Tasmanian
material is 3.46% when all bases are considered individually and 2.86% when
indels are counted asa single event. There is no standard by which two collections
can be considered different species based on ITS sequence differences but based
Cruentomycena gen. nov. for Mycena viscidocruenta ... 135
on data for within-species variation from the Great Smoky Mountains National
Park in the United States (unpub. data), a 2.86% sequence divergence usually
represents different species. Additional differences including a reduced viscid
deposition on stipe and pileus, only one type of cheilocystidia, slightly wider
spores and the specific substrate preference of Alnus catkins in Kedrovaya
suggests that the Kedrovayan collections may have diverged sufficiently to be
separate taxonomic entities. Cleland noted that basidiomata of C. viscidocruenta
were viscid when wet, implying that the viscid nature of C. viscidocruenta from
Australia may be a function of environmental conditions but Kedrovayan
collections were made in very wet conditions and still lacked the trait.
Acknowledgments
The authors extend thanks to curators at K and AD for expediting herbarium loans of
M. viscidocruenta. Administration of the Far Eastern Branch, of the Russian Academy
of Sciences, Vladivostok, arranged use of facilities at Kedrovaya Pad Nature Reserve, and
we offer our thanks. Collections from the Great Smoky Mountains National Park were
supported by NSF DEB-0338699. Thanks are due to Drs. Andrew S. Methven and D.
Jean Lodge for pre-submission reviews.
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Volume 105, pp. 137-147 July-September 2008
Caribbean heterobasidiomycetes:
3. British Virgin Islands
PETER ROBERTS
p.roberts@rbgkew.org.uk
Mycology Section, Royal Botanic Gardens
Kew, Surrey TW9 3AB, UK
Abstract — Seventeen species of heterobasidioid fungi belonging to the orders
Auriculariales, Ceratobasidiales, Dacrymycetales, Exidiales, Tremellales, and Tulasnellales
are described or reported from the British Virgin Islands. All are newly recorded from
the territory. Endoperplexa tortola and Heterochaete inconspicua are described as new
species. Heterochaete tenuicula and Serendipita vermifera are newly recorded from the
Americas. Ceratobasidium calosporum is newly recorded from the Caribbean area.
Keywords — cloud forest, xerophytic forest, Basidiomycetes
The species recorded in this paper were collected by the author and colleagues in
October 2001 during a brief visit to the islands of Guana (18°29’N, 63°33’40” W)
and Tortola (Sage Mountain area, 18°28’N, 64°33°40” W) in the British Overseas
Territory of the Virgin Islands. Vegetation on the former island, dry at the time
of visit, is mainly xerophytic scrub and semi-natural deciduous forest, rising
from the coast up to 246 m. The island is not permanently inhabited and has
been the object of extensive biodiversity surveys (Lazell 2005). Vegetation in the
Sage Mountain area of Tortola (near, but outside the National Park) is mainly
secondary cloud forest, at around 500 m elevation, with a mix of deciduous
trees.
Two previous papers in this series have dealt with species from the Dominican
Republic (Roberts 2003b) and Jamaica (Roberts 2006). Apart from two
unnamed Auricularia and Heterochaete species listed from Guana Island in
the checklist of Caribbean fungi by Minter et al. (2001) and an initial list of
Guana fungi in Lodge & Roberts (2005), no previous published references to
heterobasidiomycetes from the British Virgin Islands are known.
138 ... Roberts
Auriculariales
Auricularia cornea Ehrenb., in Nees von Esenbeck, Horae Phys. Berol.: 91 (1820)
= Auricularia polytricha (Mont.) Sacc., in Berlese & Saccardo, Atti. R. Inst. Veneto VI.
3: 722 (1885)
SPECIMENS EXAMINED — GUANA: NORTH Bay, on partly fallen branch, 4 Oct. 2001, D,J.
Lodge (Roberts GA803), K(M) 90569; TORTOLA: SAGE MOUNTAIN, TRAIL TO NATIONAL
PARK, on log, 6 Oct. 2001, Roberts GA830, K(M) 90571.
Auricularia cornea is a common and conspicuous pantropical species,
distinguished by its smooth hymenium and hirsute, but unzoned pileus. In the
present series of papers, the species has also been recorded from the Dominican
Republic and Jamaica (Roberts 2003b, 2006).
Ceratobasidiales
Ceratobasidium calosporum D.P. Rogers,
Univ. Iowa Stud. Nat. Hist. 17: 5 (1935) Fic. 1
SPECIMEN EXAMINED — TORTOLA: SAGE MOUNTAIN, TRAIL TO NATIONAL PARK, in
hymenium of Botryobasidium sp. on fallen branch, 6 Oct. 2001, Roberts GA869, K(M)
132766.
Ceratobasidium calosporum is distinguished by its bisterigmate basidia,
clampless hyphae, and vermiform basidiospores. Though it sometimes forms
thin, effused basidiomes, it can also occur more sparsely and, as in the present
collection, consist of just a few basidia-bearing hyphae. The species was
originally described from North America and has not previously been reported
from the Caribbean area.
Dacrymycetales
Dacryopinax spathularia (Schwein.) G.W. Martin, Lloydia 11: 116 (1948)
SPECIMEN EXAMINED — TORTOLA: SAGE MOUNTAIN, TRAIL TO NATIONAL PARK, on
fallen branch, 6 Oct. 2001, Roberts GA835, K(M) 117617.
This isa ubiquitous warm temperate and tropical species, producing conspicuous,
spathulate basidiomes with single-septate basidiospores. In the present series of
papers, the species has also been recorded from the Dominican Republic and
Jamaica (Roberts 2003b, 2006).
Exidiales
Basidiodendron cinereum (Bres.) Luck-Allen, Can. J. Bot. 41: 1043 (1963)
SPECIMENS EXAMINED — TORTOLA: SAGE MOUNTAIN, TRAIL TO NATIONAL PARK, on
branch, 6 Oct. 2001, Roberts GA858, K(M) 133082; same details, Roberts GA924, K(M)
132940
Caribbean heterobasidiomycetes (Virgin Islands) ... 139
Fic. 1. Ceratobasidium calosporum (Tortola, K(M) 132766).
Vermiform basidiospores; mature and immature, bisterigmate basidia
arising from clampless hyphae.
Basidiodendron cinereum is a cosmopolitan species or species complex,
normally producing thin, effused basidiomes with gloeocystidia and oblong
basidiospores. In the present series of papers, the species has also been recorded
from Jamaica (Roberts 2006).
The collection GA858 (cited above) is atypical in producing a thick, crust-
like basidiome dotted with small, white tubercules. Microscopically, it has
basidiospores and gloeocystidia typical of B. cinereum, but the hymenium
also has abundant mineral matter throughout. It may well represent a distinct
taxon.
Basidiodendron radians (Rick) P. Roberts, Kew Bull. 56: 170 (2001)
SPECIMEN EXAMINED — TORTOLA: SAGE MOUNTAIN, TRAIL TO NATIONAL PARK, on
stick, 6 Oct. 2001, Roberts GA895, K(M) 117625.
Basidiodendron radians is similar to B. cinereum (above), but produces
subglobose to ellipsoid basidiospores. The species was originally described
from Brazil and appears to be cosmopolitan. In the present series of papers, the
species has previously been recorded from Jamaica (Roberts 2006).
140 ... Roberts
Basidiodendron spinosum (L.S. Olive) Wojewoda,
Mala Flora Grzybow II: 91 (1981)
SPECIMEN EXAMINED — TORTOLA: SAGE MOUNTAIN, TRAIL TO NATIONAL PARK, on
fallen wood, 8 Oct. 2001, Roberts GA908, K(M) 121613.
Basidiodendron spinosum is a cosmopolitan species, recognized microscopically
by its distinctly warted basidiospores. It was originally described from Tahiti
and in the present series of papers has previously been recorded from Jamaica
(Roberts 2006).
Eichleriella leveilleana (Berk. & M.A. Curtis) Burt,
Ann. Missouri Bot. Gard. 2: 744 (1915)
SPECIMEN EXAMINED — GUANA: IGUANA HEAD, LEEWARD TRAIL, on fallen sticks, 5 Oct.
2001, Roberts GA815, K(M) 90647.
The species forms reddish, effused, stereoid basidiomes. It was originally
described from North America and in the present series of papers has previously
been reported from Jamaica (Roberts 2006).
Endoperplexa tortola P. Roberts sp. nov. Fic. 2
MycoBank 511665
Basidiomata effusa, laevia, ceracea, pallide caesiogrisea. Systema hypharum monomiticum.
Hyphae agglutinatae, 1-2 um latae, fibulatae. Hyphidia non visa. Cystidia desunt. Basidia
tremelloidea, 2-4-septata, globosa vel ovoidea, 8-11.5 x 7-10.5 um. Basidiosporae
oblongae, 7-8.5 x 3.5-4 um.
Ho.LotyPeE — BRITISH VIRGIN ISLANDS. ToRTOLA: SAGE MOUNTAIN, TRAIL TO
NATIONAL PARK, on dead woody stems, 6 Oct. 2001, Roberts GA892, K(M) 132962.
MACROCHARACTERS — BASIDIOMES effused, smooth, ceraceous, thin, in small
patches without distinct margins, pale blue-grey drying to a thin but visible
blue-grey film.
MICROCHARACTERS — HYPHAL SYSTEM monomitic; HYPHAE hyaline,
agglutinated and difficult to see clearly in dried material, c. 1-2 um wide, thin-
walled, with clamp-connexions; HYPHIDIA not seen and apparently absent;
CYSTIDIA absent; BASIDIA tremelloid, 2-4-celled, globose to ovoid, 8-11.5 x
7-10.5 tum, not or scarcely stalked; sTERIGMATA short; BASIDIOSPORES oblong
(Q = 1.8-2.1), 7-8.5 x 3.5-4 um.
The genus Endoperplexa P. Roberts was introduced for thin, effused, exidioid
species having agglutinated hyphae, small basidia with short sterigmata, hyaline
cystidia, and no hyphidial layer (Roberts 1993). Endoperplexa tortola matches _
these criteria well, except for the apparent lack of cystidia. No more appropriate
genus suggests itself.
The lack of cystidia distinguishes E. tortola from all other Endoperplexa species.
The European E. subfarinacea (Hauerslev) P. Roberts additionally has rather
Caribbean heterobasidiomycetes (Virgin Islands) ... 141
10 pm
Fic. 2. Endoperplexa tortola (Holotype, K(M) 132962).
Oblong basidiospores;
mature and immature basidia arising from agglutinated, thin-walled hyphae.
more cylindrical basidiospores, whilst E. phlebioides P. Roberts, recently
described from Belize, has basidiospores which are oblong but distinctly smaller.
Endoperplexa obscura (G.W. Martin) P. Roberts, described from Panama, has
basidiospores which are oblong and of similar size, but are subfusiform.
Exidiopsis galzinii (Bres.) Killerm., in Engler & Prantl,
Die nattirlichen Planzenfamilien (2nd ed.) 6: 113 (1928)
SPECIMEN EXAMINED — TORTOLA: SAGE MOUNTAIN, TRAIL NEAR NATIONAL PARK, on
fallen decorticated wood, 8 Oct. 2001, Roberts GA910, K(M) 117750.
Exidiopsis galzinii is distinguished by its effused, ceraceous-gelatinous
basidiomes (grey when fresh, in the collection cited above), its abundant
enclosed gloeocystidia, layer of branched hyphidia, and myxarioid (stalked)
basidia. The species was originally described from France, but appears to be
cosmopolitan. It was redescribed and illustrated from the Dominican Republic
by Roberts (2003b).
Heterochaete crassa Bodman, Mycologia 41: 531 (1949)
SPECIMEN EXAMINED — TORTOLA: SAGE MOUNTAIN, TRAIL TO NATIONAL PARK, on
stick, 6 Oct. 2001, Roberts GA873, K(M) 116906
Heterochaete crassa has eftused, pinkish-brown basidiomes with abundant
hyphal pegs, a dimitic hyphal system with skeletal hyphae predominating, and
142 ... Roberts
Fic. 3. Heterochaete inconspicua (Holotype, K(M) 132950).
Cylindrical basidiospores;
mature and immature basidia with an irregularly fusiform cystidium.
rather narrow, clavate basidia. It was originally described from North America
and was redescribed from Jamaica and illustrated by Roberts (2006).
Heterochaete inconspicua P. Roberts sp. nov. Fic. 3
MycoBank 511664
Basidiomata effusa, ceracea, tenuissima, grisea, in sicco ochracea; setulis irregularis,
albidis, brevis (c. 30-60 um), c. 8-10 per mm. Systema hypharum monomiticum. Hyphae
hyalinae, 2-3 um latae, tenuitunicatae, fibulatae. Cystidia sparsa, clavata vel fusoidea,
usque ad 30 x 8 um, tenuitunicata, hyalina. Basidia tremelloidea, 4-septata, subglobosa
vel ellipsoidea, 10-12 x 8-10 um. Basidiosporae cylindraceae, 9-11 x 3.5-4 um,
suballantoideae vel allantoideae.
Caribbean heterobasidiomycetes (Virgin Islands) ... 143
HoLotyPE — BRITISH VIRGIN ISLANDS. TorToLa: SAGE MOUNTAIN, TRAIL TO
NATIONAL PARK, on dead woody stems, 6 Oct. 2001, Roberts GA850, K(M) 132950.
MACROCHARACTERS — BASIDIOMES effused, adnate, ceraceous, very thin (c.
40-50 um thick) and visible only as greyish patches, ochraceous when dried,
appearing grandinioid under lens and covered in fine, whitish, irregular and
often coalescing pegs or spicules, c. 30-60 tm long, c. 8-10 per mm. Margins
not distinct. Spicules sometimes tubular and discrete, sometimes amalgamating,
mainly composed of agglomerated mineral matter, but interspersed with some
thin-walled hyphae.
MICROCHARACTERS — HYPHAL SYSTEM monomitic; HYPHAE agglutinated
in a partly gelatinized matrix, hyaline, 2-3 um wide, with thin or slightly
refractive walls, with clamp-connexions; HYPHIDIA branched, but not always
seen and may break up into mineral particles at maturity; cysTIDIA scattered
and not always present, irregularly clavate to fusiform, up to 30 x 8 um, thin-
walled, hyaline or with pale yellowish contents; BAsIDIA tremelloid, septate, 4-
spored, subglobose to ellipsoid, 10-12 x 8-10 um, not stalked; sTERIGMATA
sinuous, elongated; BASIDIOSPORES cylindrical (Q = 2.2-3.0), 9-11 x 3.5-4 um,
suballantoid to allantoid.
Heterochaete inconspicua is distinguished macroscopically by its thin, ceraceous
basidiomes with irregularly scattered, often coalescing, small pegs or spicules
and microscopically by the mainly mineral composition of these spicules, the
comparatively small basidia, and the small, narrow basidiospores. When dried,
the colour and grandinioid appearance may suggest a species of Basidiodendron
Rick. The scattered cystidia are infrequent and may be absent in some mounts.
Heterochaete verruculosa (M6ller) Bodman is a similarly thin, ceraceous
species, but differs inter alia in its more regular hyphal pegs which are
not mineralized. Heterochaete sanctae-martae Bodman (redescribed and
illustrated in Roberts 2003) is also thin and ceraceous, but has distinctly wider
basidiospores (Q = 1.8-2.2). Both these species lack cystidia.
Heterochaete shearii (Burt) Burt, Ann. Missouri Bot. Gard. 8: 377 (1921)
SPECIMEN EXAMINED —TORTOLA: SAGE MOUNTAIN, TRAIL TO NATIONAL PARK, on
partly fallen stick, 6 Oct. 2001, Roberts GA846, K(M) 90583.
Heterochaete shearii can be distinguished macroscopically by its effused, adnate,
greyish-brown basidiomes, bruising brown, with abundant but irregular,
tubular hyphal pegs, and microscopically by its brown subicular hyphae,
frequent cystidia-like elements in the hymenium, and consistently bisterigmate
basidia (illustrated in Roberts 2001). The species, originally described from
North America, is common and widespread. In the present series of papers, it
has previously been recorded from Jamaica (Roberts 2006).
144 ... Roberts
Fic. 4. Heterochaete tenuicula (Guana, K(M) 90575). Cylindrical basidiospores; section of
hymenium showing mature and immature, bisterigmate basidia, an irregularly cylindrical
cystidium, and branched, sometimes thick-walled hyphidia; thick-walled skeletal hyphae.
Heterochaete tenuicula (Lév.) Pat., Bull. Soc. Mycol. Fr. 10: 76 (1894) Fi. 4-
= Hydnum tenuiculum Lév., Ann. Sci. Nat., Bot., sér. 3, 5: 145 (1846)
= Hydnum delicatum Klotzsch ex Berk. [nom. illeg., non H. delicatum Schwein., 1832], Ann. Mag.
Nat. Hist. 3: 395 (1839)
= Heterochaete delicata Bres., Hedwigia 53: 77 (1912)
Caribbean heterobasidiomycetes (Virgin Islands) ... 145
MACROCHARACTERS — BASIDIOMES effused, typically arising as circular patches
which subsequently coalesce; papery and dry, cream to pale buff when fresh,
pale buff when dried, with 5-8, erect, narrow, tubular, hyphal pegs per mm, the
margins distinct, smooth, velvety, often reflexed.
MICROCHARACTERS — HYPHAL SYSTEM dimitic; HYPHAE in hymenium hyaline,
1.5-3 um wide, thin- to thick-walled, with clamp-connexions; skeletal hyphae
ochre-brown, 2-3.5 ttm wide, thick-walled. Hyphal pegs 100-250 x 40-80 um,
arising from below the hymenial layer and mainly composed of brown skeletal
hyphae; HYPHIDIA variously nodulose to branched, thin- to thick-walled;
CYSTIDIA present in the hymenium as occasional clavate, cylindrical, fusiform,
or irregular elements; BAsIDIA tremelloid, 2-celled, oblong, 15-17 x 9-10.5
[tm; STERIGMATA elongated, sinuous; BASIDIOSPORES cylindrical (Q = 3.1-3.6),
some ventrally depressed, 16-21 x 5.5-6 ttm, germination not seen.
SPECIMENS EXAMINED — GUANA: IGUANA HEAD, on partly fallen twig, 5 Oct. 2001,
Roberts GA816, K(M) 90670; Nort Bay, on partly fallen twigs, 9 Oct. 2001, Roberts
GA943, K(M) 90575.
Heterochaete tenuicula is a common, pantropical species, recognizable in the
field by its effused, papery, buff basidiomes with distinct, often reflexed margins,
and microscopically by its two-spored basidia and brown, thick-walled, skeletal
hyphae. The spores are considerably larger than those of the equally bisterigmate
H. shearii (above). The species was originally described from Java and has not
previously been reported from the Americas.
Serendipita vermifera (Oberw.) P. Roberts, Mycol. Res. 97: 474 (1993) Fi@u5
SPECIMEN EXAMINED — TORTOLA: SAGE MOUNTAIN, TRAIL TO NATIONAL PARK, in
hymenium of Botryobasidium sp. on fallen branch, 6 Oct. 2001, Roberts GA875, K(M)
132767.
Serendipita vermifera typically forms no visible basidiomes, but is found in
the hymenia of corticioid fungi (particularly Botryobasidium species) where
it produces clusters of tremelloid basidia and vermiform basidiospores. The
species was originally described from Germany and is widespread, though
seldom collected, in Europe. It has not previously been reported from the
Americas.
Tremellales
Tremella foliacea Pers., Obs. Mycol. 2: 98 (1800)
SPECIMEN EXAMINED — TORTOLA: SAGE MOUNTAIN, TRAIL TO NATIONAL PARK, on
half-fallen stick, 6 Oct. 2001, Roberts GA834, K(M) 132939.
Tremella foliacea is a cosmopolitan species with brown, typically foliaceous
basidiomes (though small and more lobate in the collection cited above). In
the present series of papers, the species has been reported from the Dominican
146 ... Roberts
Fic. 5. Serendipita vermifera (Tortola, K(M) 132767).
Vermiform basidiospores; mature and immature basidia.
Republic (Roberts 2003b). Specimens from England were described and
illustrated in Roberts (1999) and from Brazil in Roberts & de Meijer (1997). It is
normally assocated with the mycelia of Stereum species, though no basidiomes
of the latter were seen in the Tortola collection.
Tremella fuciformis Berk., Hooker’s J. Bot. Kew Gard. Misc. 8: 277 (1856)
SPECIMEN EXAMINED — TORTOLA: SAGE MOUNTAIN, TRAIL TO NATIONAL PARK, On dead
standing pole, 6 Oct. 2001, Roberts GA832, K(M) 90275.
Tremella fuciformis is a common, pantropical species, originally described
from Brazil and distinguished by its delicate, white, fuciform basidiomes, small
basidia and basidiospores (illustrated in Roberts & de Meijer 1997). In the
present series of papers, the species has been reported from Jamaica (Roberts
2006).
Tulasnellales
Tulasnella eichleriana Bres., Ann. Mycol. 1: 113 (1903)
SPECIMENS EXAMINED — TORTOLA: SAGE MOUNTAIN, TRAIL NEAR NATIONAL PARK, On
fallen wood, 8 Oct. 2001, Roberts GA917, K(M) 117623; same details, Roberts GA940,
K(M) 117619.
Caribbean heterobasidiomycetes (Virgin Islands) ... 147
Tulasnella eichleriana is a cosmopolitan species or species complex, originally
described from Poland and redescribed and illustrated by Roberts (1994). The
species can be recognized by its effused, ceraceous, grey-violaceous basidiomes,
clampless hyphae, and small, subglobose to ellipsoid basidiospores. Tulasnella
eichleriana has been recorded in the Caribbean from the Dominican Republic
and Jamaica (Roberts 2003b, 2006).
Acknowledgments
Thanks to Dr Karen Nakasone and Dr D. Jean Lodge for refereeing the paper. Fieldwork
and travel within the British Virgin Islands were supported and facilitated by The
Conservation Agency (Rhode Island, USA). Thanks go to Dr J. Lazell, president of The
Conservation Agency, and to Dr D. Jean Lodge, Center for Forest Mycology Research
(Puerto Rico), for helping organize the visit.
References
Lazell J. 2005. Island. Fact and theory in nature. Berkeley, Univ. California Press.
Lodge DJ, Roberts P. 2005. Fungi. In Lazell J., Island. Fact and theory in nature. 171-174. Berkeley,
Univ. California Press.
Minter DW, Rodriguez Hernandez M, Mena Portales J. 2001. Fungi of the Caribbean: an annotated
checklist. Middlesex, PDMS.
Roberts P. 1993. Exidiopsis species from Devon, including the new segregate genera Ceratosebacina,
Endoperplexa, Microsebacina, and Serendipita. Mycol. Res. 97: 467-478.
Roberts P. 1994. Globose and ellipsoid-spored Tulasnella species from Devon and Surrey, with a
key to the genus in Europe. Mycol. Res. 98: 1431-1452.
Roberts P. 1999. British Tremella species II. Tremella encephala, T. steidleri, and T. foliacea.
Mycologist 13: 127-131.
Roberts P. 2001. Heterobasidiomycetes from Korup National Park, Cameroon. Kew Bull. 56: 163-
187.
Roberts P. 2003. Heterobasidiomycetes from Rancho Grande, Venezuela. Mycotaxon 87: 25-41.
Roberts P. 2003b. Caribbean heterobasidiomycetes. 1. Dominican Republic. Mycotaxon 87: 187-
201.
Roberts P. 2006. Caribbean heterobasidiomycetes. 2. Jamaica. Mycotaxon 96: 83-107.
Roberts P, de Meijer AAR. 1997. Macromycetes from the state of Parana, Brazil 6: Sirobasidiaceae
& Tremellaceae. Mycotaxon 64: 261-283.
MYCOTAXON
Volume 105, pp. 149-164 July-September 2008
A study of lichenicolous species of Polysporina
(Acarosporaceae)
KERRY KNUDSEN! & JANA KOCOURKOVA ”
kk999@msn.com & jana_kocourkova@nm.cz
'The Herbarium, Dept. of Botany and Plants Sciences
University of California Riverside, Ca., USA 92521-0124
*Mycological Department, National Museum
Vaclavské nam. 68, 115 79 Praha 1, Czech Republic
Abstract — Three lichenicolous species of Polysporina are recognized: Polysporina
arenacea, P. pusilla, and P. subfuscescens. Polysporina lapponica is found to be a
misapplied name to P. subfuscescens. The new combinations P. arenacea, P. subfuscescens,
and Sarcogyne lapponica are made. Sarcogyne lapponica is reported from Norway,
Sweden and Switzerland. A neotype is selected for Sarcogyne dubia. Polysporina pusilla
is reported new for Asia (Iran, Turkey) and North America (western North America).
Key words — accession fissure, algicolous, calciphiles, carbonization, species concepts
Introduction
Polysporina (Acarosporaceae) is a small and common crustose genus occurring
on acid and calcareous rock. As currently circumscribed, the genus contains 12
valid binominals (MycoBank 2008). The type species for the genus is Polysporina
simplex (Davies) Vézda. Polysporina occurs in both the northern and southern
hemispheres (Knudsen 2008a). The most recently described species was
Polysporina frigida Kantvilas & Seppelt (Kantvilas & Seppelt 2006).
The main characters of Polysporina are a non-amyloid tholus (Acarospora-
type ascus), polyspory (50-200 ascospores per ascus), a non-thalline margin,
a carbonized exciple, and the build-up of carbonized accretions on the surface
of the apothecial disc (Vézda 1978, Golubkova 1988, Kantvilas 1998, Knudsen
2008a). Vézda (1978) used the character of branching and anastomosing
paraphyses in P. simplex to segregate the genus Polysporina from Sarcogyne.
Based on our observations, branching and anastomosing paraphyses may be
frequent or infrequent in the same species. Branching of paraphyses can be
observed in many Sarcogyne species and some Acarospora species, particularly
in A. smaragdula (Wahlenb.) A. Massal. The width of the apices of paraphyses
also varied in our observations.
150 ... Knudsen & Kocourkova
Carbonization is the build-up of pigmentation in fungal structures. In
Acarosporaceae, the most extreme carbonization of the exciple is seen in species
Polysporina and Sarcogyne clavus (DC.) Kremp., with complete death and fusion
of the excipular hyphae. Polysporina is very close to the core group of Sarcogyne,
e.g. S. clavus and S. privigna (Ach.) A. Massal., both of which have carbonized
exciples. Polysporina species differ morphologically from Sarcogyne s. str. only
in the build-up of carbonized accretions (umbos and ridges) on the apothecial
surface that gives the genus its characteristic look. In early development
of P. urceolata (Anzi) Brodo, the apothecia do not usually have carbonized
accretions on the disc surface, but in other species the accreted structures are
usually apparent at all stages of development of apothecia. The development
of carbonization on the surface of the epihymenium, the formation of sterile
columns of hyphae in the hymenium, and the vegetative division of apothecia
all need further study.
Polysporina simplex has a basal position in the phylogeny of Acarosporaceae
(Reeb et al. 2004) but more molecular analysis of Polysporina species is necessary
because of problems with other specimens sequenced (Reeb, pers. comm.). The
differing placement of P simplex in the phylogeny proposed by Crewe et al.
(2006) was due to the mis-determination of a specimen of Sarcogyne clavus as
P. simplex (Martin Westberg, pers. comm.).
In Polysporina the species concepts are primarily based on size of apothecia,
height of hymenium, paraphyses width, ascospore size and form, and substrate.
The taxonomy of Polysporina is derived from the work of Magnusson (1935).
Magnusson’s species concepts are generally very narrow (Knudsen 2004) though
occasionally as in the Acarospora smaragdula group his suite of characters led
him to overlook other characters such as differences in thallus macrostructure
(Knudsen 2008b).
This study began in a sunny wash in the southwestern Mojave Desert in
California while we were out collecting lichenicolous fungi. The taxon currently
known as Polysporina lapponica (Triebel et. al. 1991; Knudsen 2005, 2008a) is
common in the Mojave Desert (Knudsen & La Doux 2005, 2006; Knudsen &
Werth 2008). We wondered whether the taxon in California was the same as
the taxon in the Czech Republic because the southern California specimens
usually have wider (usually 2-2.5 um) ascospores and wider (2 um) paraphyses
than the average Czech specimens with narrower (usually 1.5 um) ascospores
and narrower (1-1.5 um) paraphyses. The ascomata of California specimens
appeared larger and more superficial too.
To decide whether specimens from Europe and North America were
conspecific or distinct species, we had to revise Magnusson’s taxonomy. We
began with an investigation of the Acharius type of Lecidea lapponica and then
examined lichenicolous specimens from Europe, Asia, and North America.
A study of Polysporina ... 151
Methods and materials
Sections were prepared by hand and examined in water, lactophenol cotton
blue, 10% KOH, and I (Lugol). Reactions were tested in I and I with pre-
treatment with KOH (K/I). Ascospore measurements were made in water and
measurements of apothecial structures in water and KOH. Values in parenthesis
represent lowest or highest measurements. A new term accession fissure is used
to describe the separation of apothecia during vegetative division.
Specimens were studied from ASU, B, COLO, FR, GZU, H, LE, LD, MIN,
NY, PRM, S, UCR, UPS, and W.
Introductory discussion
Lichenicolous Polysporina are non-lichenized fungi with both endolithic and
endokapylic thalli. The mycelium of the endolithic thallus occurs in the upper
1 mm of rock. It is either independent of a host, migratory from one host to
another (observable as a line of apothecia between hosts), or is parasitic on the
endolithic thalli of other lichens like Lecidea laboriosa Mill. Arg. and probably
algicolous especially in the case of P pusilla. The thallus when endokapylic
may enter the host either through ascospores lodging on thallus or through
infiltration of the host from below by the endolithic thallus.
All three taxa discussed here are not host specific, though they may be more
common on one host than another in particular biotas. All three species are
pathogenic, degrading the fungal host and exhausting the algal layer. Collector
bias is for specimens where the host is usually intact during the early stage of
infection. These specimens can also give the impression that the fungus may be
a parasymbiont. But even in these specimens ascomata production has usually
already been suppressed in the host.
The lichenicolous species
1. Polysporina subfuscescens (Nyl.) K. Knudsen & Kocourk., comb. nov.
MycoBAnk MB 511741
Type: FRANCE. CoLLiurE, 1872 (H-NYL 24518, HOLOTYPE).
Basionym. Lecanora subfuscescens Nyl., Flora (Regensburg) 56: 199 (1873).
= Biatorella subfuscescens (Nyl.) H. Olivier, Bull. Acad. Intern. Geogr. Bot 11: 56
(1902).
= Acarospora subfuscescens (Nyl.) H. Magn., Rabh. Krypt.-FL., ed. 2, 9(5:1): 146
(1935).
= Sarcogyne subfuscescens (Nyl.) Boistel, Nouv. Fl. Lichens 2: 228 (1903).
= Acarospora silesiaca H. Magn., Rabh. Krypt.-Fl., ed. 2, 9(5:1): 145 (1935). Type:
POLAND. Dolnoslaskie. swiDNIcA, 1890, Eitner (UPS, HOLOTYPE).
= Acarospora sordida var. urbana H. Magn., Kungl.-Akad. Handlingar ser. 3, 7(4): 126
(1929). Type: SLOVAKIA. BRATISLAVA, ST. GEORGEN (SVATY JUR), on a stone bridge
in vineyard, 21 VII 1922, Magnusson 6868b (UPS, HOLOTYPE; S, ISOTYPE).
152 ... Knudsen & Kocourkova
= Acarospora tyroliensis H. Magn., Rabh. Krypt.-FL., ed. 2, 9(5:1): 149 (1935). Type:
AUSTRIA. TYROL, ALPS, hb. Zwackh (H-NYL 24560, HOLOTYPE).
= Sarcogyne scabra Nyl., Lich. Japon., p. 49 not. (1890). Type: AUSTRIA. TYROL, ALPS,
hb. Zwackh (H-NYL 24560, HOLOTYPE).
= Polysporina ferruginea (Lettau) M. Steiner ex Kantvilas, Lichenologist 30(6): 557
(1998). Type: not seen, see discussion.
= Sarcogyne simplex. f. ferruginea Lettau, Feddes Repert. Spec. Nov. Regni Veg.,
57: 73 (1955).
= Sarcogyne bicolor H. Magn. Ann. Crypt. Exot., 7: 130-131 (1934). Type: USA.
CALIFORNIA: Los Angeles Co. SANTA MONICA MOUNTAINS, OCEAN BLUFFS, POINT
DUME, VIII 1928, Hasse (FH, HOLOTYPE).
= Acarospora gyrosa N. S. Golubk., Novitates Systematicae Plantarum non Vascularium
9: 220 (1972). Type: TADZHIKISTAN. vosTOCHNY] PAMIR, MURGAB, CHECHEKTA
R., 3860 m, on silicate boulder, 29 VII 1964, Golubkova (LE 378, HOLOTYPE).
Sarcogyne dubia H. Magn., Rabh. Krypt.-Fl., ed. 2, 9(5:1): 72. Type: FRANCE.
HERAULT, L’AGDE, (sur) laves, 1935, hb. J. Maheu (UPS, NEoTYPE designated
here).
= Polysporina sinensis (H. Magn.) N. S. Golubk., Konspekt Flory Lishainikov
Mongol'’skoi Naradnoi Respubliki (Leningrad). p. 138 (1981). Type: MONGOLIA.
KANSU, HEI-KU (SHARGALTEIN), 3 IV 1932, Bohlin 85a (S, HOLOTYPE).
= Sarcogyne sinensis H. Magn., Lich. from Central Asia, p. 70 (1940).
DEscCRIPTION — Lichenicolous, THALLUS endokapylic or endolithic, vegetative
hyphae hyaline, usually less than 1 mm diam, I-.
APOTHECIA immersed to superficial, dull black or brownish black when dry,
always black when wet, round and convex to irregular, 0.3-1.0(-3.0) mm diam,
usually less than 0.5 mm tall, dispersed or clustered; larger apothecia, between
1-3 mm, usually in a process of division, forming agglomerations with deep
accession fissures, initially forming angular edges when division completed;
disc black with accretions of carbonization in irregular structures 50-150 um
tall, with small patches of reddish smooth un-carbonized disc surface; margin
fissured, rough. EXCIPULUM of narrow hyphae to 100 um thick, especially near
surface of disc, outer layer carbonized, inner layer narrow, yellowish to hyaline.
HYMENIUM (80-)100-140(-180) um tall, hyaline to yellowish-orange, often
divided by sterile columns of conglutinated hyphae, usually I+ blue eventually
turning red but often reaction various from red to yellow-green, epihymenium
10-15 um thick, conglutinated in a dark brownish or reddish-brown gel.
PARAPHYSES coherent, 1-2 (-3) um diam at mid-level, sometimes various in
size in same hymenium, frequently or rarely branching, sometimes thicker
where branched or rarely paraphyses 2-3 um diam mixed with narrower
paraphyses, with or without oil drops, septate, rarely constricted at septa, cells
4-7 um long, usually becoming shorter in upper third, apices barely expanded
to 3 um wide. asci clavate to subcylindrical, usually 50-110 x 12-20 um,
100 to 200 ascospores per ascus, varying, sometimes with between 50-100
ascospores per ascus. ASCOSPORES (3.0—)4.0-4.5(-5.0) x (1.0—)1.5-2.0(-3.0)
A study of Polysporina ... 153
um, narrowly to broadly ellipsoid. The spores varied in size, sometimes
depending on amount of ascospores per ascus, both between specimens and
even in same specimen. SUBHYMENIUM 20-50 um thick, usually hyaline,
sometimes reddish to golden. HYPOTHECIUM (20-)30-45 um thick, of narrow
hyaline prosoplectenchyma, ca. 0.5-1.0 um diam, forming often uneven border
in thallus of host, confluent with vegetative hyphae, and often appearing much
deeper but actually indistinguishable from vegetative hyphae infiltrating host
from below. CONIDIOMATA not observed.
ECOLOGY AND SUBSTRATE - Appearing to prefer sunny open habitats, on
various crustose hosts including species of Acarospora, Buellia, Caloplaca,
Candelariella, and Lecidea on usually siliceous rock.
DISTRIBUTION — North America, Europe, Asia, in the northern hemisphere, up
to 1869 m.
Discussion - Polysporina subfuscescens is a widely distributed and successful
parasitic fungus. It is an aggressive parasite, suppressing ascomata production
of host, and eventually depleting algal layer and destroying the host, dissolving
the stratified lichen structure and reducing it to a clumps of ecorticate white
gelatinized hyphae. Then the ascomata of the parasite eventually atrophy.
Polysporina subfuscescens infects the host in two ways. Firstly, the ascospores
can lodge on the host. The excipulum of the ascomata in an early stage of
infection forms a distinct and narrow hypothecium in the thallus of host and
vegetative hyphae are often scant. Secondly, the hyphae of P. subfuscescens can
infiltrate the thallus of host from below forming an extended structure beneath
the apothecium that can be mistaken for a deep hypothecium.
The endolithic thallus of Polysporina subfuscescens may be partially lichenized
as a few lichenized algal cells are sometimes found in rock substrate or these
may be remnants of a parasitized endolithic thallus of a saxicolous lichen.
Though our investigation began with the hypothesis that there were two
taxa, the observed variation was wider in ascomata size, height of hymenium,
paraphyses width, and ascospore size than in earlier descriptions (Magnusson
1935, Vézda 1970, Kantvilas 1998). The branching of paraphyses was also varied
from frequent to infrequent. We found no apparent regional distribution of
any variation either. Thus we agree with Triebel’s broad concept of the taxon
(Triebel et al. 1991).
Since the name Polysporina lapponica was misapplied to this taxon (see
Excluded Species below), the earliest name we found for this taxon was Lecanora
subfuscescens and a new combination in Polysporina is proposed.
The synonyms proposed were generally treated by the authors as different
species or infraspecifc taxa based on the misconception that the thallus of host
is the thallus of the lichenicolous fungus.
154 ... Knudsen & Kocourkova
The exception was that Sarcogyne dubia (Magnusson 1935) was recognized
as a lichenicolous fungus. The original circumscription of the species differs
completely from that of Vézda (1970). Based by Magnusson (1935) on a
single collection from South France collected by de Crozals in the herbarium
of Bouly de Lesdain, it differed in having fifty ascospores per ascus, all 3 um
wide, and with no description of branching paraphyses. This specimen was
apparently originally included in Acarospora lapponica (Ach. ex Schaer.) Th.
Fr. (Magnusson 1929). The holotype could not be located and is presumed
lost during World War II. A neotype from South France is selected here. It is
actually quite similar to the variation of some specimens from California and
Bulgaria with most ascospore widths between 2-3 um and with occasional asci
with 50-100 ascospores per ascus.
Using the narrow concepts inherited from Magnusson, Kantvilas (1998)
recognized Polysporina ferruginea as separate from his concept of P lapponica
but specimens examined by us did not differ from other specimens of P
subfuscescens. We are unsure of the status of the holotype, but base our
synonymization on an excellent paratype collected by Lettau and annotated by
Steiner from B.
The holotype of Polysporina sinensis is definitely a lichenicolous fungus
occurring on the thallus of a crustose lichen (see drawing in Golubkova
1988). The holotype was too scant to dissect. Magnusson writes “Spores at
least 50, 5-7 x 3 um.” We are skeptical of Magnusson’s measurements based
on our experience of examining his species described from specimens from
expeditions and include the name as synonym of P. subfuscescens based on
other specimens seen from Asia. Other specimens determined by Magnusson
(1940) as P. sinensis, however, belong to a lichenized taxon that will be treated
in a forthcoming publication.
We did not see types on which the following names are based but assume they
are synonyms based on the literature (Magnusson 1935): Acarospora lapponica
var. silesiaca H. Magn., Acarospora sordida Wedd. [Acarospora subfuscescens
var. sordida (Wedd.) H. Magn.], Acarospora sordida var. lojkeana H. Magn.,
Lecidea simplex var. chloroclinella Wedd., Sarcogyne scabra var. canasiacensis
Hue [Sarcogyne canasiacensis (Hue) H. Magn., Biatorella canasiacensis (Hue)
H. Olivier], Sarcogyne simplex var. parasitica de Lesd., and Sarcogyne simplex
var. crustosa H. Magn. We believe Acarospora sernanderi H. Magn. is also a
synonym but both types we saw were inconclusive (S!, UPS!) Sarcogyne
sphaerospora J. Steiner is another probable synonym.
Since it is impossible to distinguish Polysporina subfuscescens and P. simplex
using apothecial and ascospore measurements, overt signs of parasitism are
used primarily to distinguish them. In difficult specimens, one must first
ascertain if the Polysporina is lichenized or not. If there is no distinct algal layer
A study of Polysporina ... 155
beneath the apothecia, scratching away the top layer of the surrounding rock
substrate and observing a well-developed layer of lichenized algae in a hyphal
matrix, is the easiest way to determine P. simplex.
SELECTED SPECIMENS EXAMINED — BULGARIA. EASTERN RODOPI MOUNTAINS, KARDZI,
KALOJANCI, PROTECTED AREA, CA. 5 KM SW OF THE VILLAGE, 41°37’N, 25°31E , 500 m,
on crustose lichen on acid vulcanite stones on north-facing slope, 15 VHI 2004, Vondrak
1999 (CBFS); SANDANSKI DISTR., IN RIVER VALLEY OF SANDANSKA BISTRICA, 400 m, on
brown crustose lichen, 29 V 1968, Vézda, Lichens Selecti Exsiccati 693 (H); CZECH
REPUBLIC. Bohemia. 300 m, on Acarospora species on sandstone, 1926, Servit (UPS);
West Bohemia, KOMOTAU, 1930, Servit (UPS); Central Bohemia. PRAHA, SUCHDOL,
PROTECTED AREA SEDLECKE ROCKS, OUTCROPS ABOVE VLTAVA RIVER, 270 m, on
Acarospora fuscata on proterozoic shale, 13 X 2002, Kocourkova (PRM 907503, 907542);
PRAHA, KLUKOVICE, PROKOPSKE VALLEY, SPICAK HILL, ABOVE FORMER MILL, on top of
steep west-facing calcareous cliff, 265 m, on Acarospora fuscata on quartz, 17 XI 1999,
Kocourkova (PRM 909497); PRAHA, MALA OHRADA, ALBRECHTUV HILL BY PROKOPSKY
BROOK, 300 m, on Acarospora fuscata on diabase, 23 IX 1999, Kocourkova (PRM 90949);
PRAHA, NOVA VES, PROKOPSKE VALLEY, HEMROVY ROCKS, on southwest-facing slope
below steppe, 265 m, on Acarospora fuscata on diabase, 6 XI 1994, Kocourkova (PRM
909499); PRAHA, REPORYJE, DALEJSKE UDOLI VALLEY, ABOVE TRUNECKUV MLYN, 265 m,
on Acarospora fuscata on diabase, 6, XI. 1994, Horakova (PRM 909500); PRAHA, MALA
CHUCHLE, BRANICKY BRIDGE, 220 m, on crustose lichen on diabase rocks, 3 IV 1988,
Hordkova (PRM 887565); DISTR. RAKOVNIK, KRIVOKLATSKO PROTECTED LANDSCAPE
AREA, BETWEEN NEZABUDICE AND ROZTOKY VILLAGES, NEZABUDICKE SKALY NATURE
RESERVE, AT NORTH MARGIN ABOVE BEROUNKA RIVER, On steep southwest-facing slope,
in thin scree oak forest, 50°01.435’N, 13°50.689°W, 390 m, on Acarospora fuscata on
schist, 16 II 2007, Kocourkova & Kocourek (PRM 896204); Eastern Bohemia. HUMPOLEC,
ABOVE THE VILLAGE OF KRASONOVA, 630 m, parasitic on Acarospora fuscata, A. sinopica
& Buellia sororia, 9 V 1968, Vézda, Lichenes Selecti Exsc. 693 as Acarospora subfuscescens
(PRM 665345, S); 9 V 1968, Vézda, Lichenes Selecti Exsc. 694 as Sarcogyne dubia (PRM
665343, H, S); Moravia. MOHELNO, IN VALLEY OF RIVER JIHLAVKA, on crustose lichen on
serpentine, 1926, Suza (UPS); DENMARK. North Jutland. skKAGEN MUNI, KATBAKKER
SW OF KANDESTEDERNE, DISTRICT 1, 15 V 1992, Svane 9034-1 (UPS); RABJERG STENE,
SW OF KANDESTEDERNE, DISTRICT 1, on crustose lichen on pebbles, 31 HI 1977, Svane
506 (COLO); GREENLAND. GODHAVEN, AT THE ARCTIC STATION, 69°15’N, 53°32’ W,
12 m, 28 VII 1952, Gelting 18736c (UPS); HUNGARY. IN VILLAGE SAJOKERESZIUR, 125
m, on roof, 5 IV 1932, F Foriss, Lichenes Bikkenses Exsic. 92, det. Magusson as
Acarospora subfuscescens (H, PRM 693682); GERMANY. Hessen. TAUNUS, NW OF
LANGTAL AT NEU ANSPACH, in thin oak forest SW-exposure, 480 m, on brown crustose
lichen on gneiss, 12 V 1993, Schéller 340 (FR); WETTERAU, 150 m, on sandstone, 8 V
1993, Schéller 302 (FR); ERZGEBIRGE, ALTENBERG, 740 m, on brown crustose lichen on
basalt, 17 VII 1928, Schade 2132 (UPS); IRAN. KHOy, airport, 38°25°16” N, 44°54’05” E,
on Caloplaca transcaspica on siliceous rock, 2 V 2007, Vondrak 5837 (CBFS); JAPAN?
1850, as Sarcogyne scabra, Herb. Zwackh (S); NORWAY. More og Romsdal. NoRDAL
HD., GRONNINGSATER, 350 m, on brown crustose lichen on stone fence, 7 IX 1947,
Magnusson 20778 (UPS); Rogaland. NORTH OF HAUGESCUND, on crustose lichen on
slate roof near road, 3 VIII 1939, Magnusson 16916 (H, S, UPS); Oppland. Lom parisH,
THE GLITTERTIND AREA, ICE-CORED MORAIN OF GRASUBREEN, 1950 m, on crustose
lichen on boulders, 11 VIII 1963, Santesson 16270 (UPS); MEXICO. Baja CALIFORNIA
NORTE: ARROYO 1 KM N OF CATAVIANA, 29°43’ N, 114°40’W, 560 m, parasitic on either
156 ... Knudsen & Kocourkova
endolithic lichen or remains of unknown host, 6 I 1989, Egan 13918 (MIN); Baya
CALIFORNIA: 3 KM WNW OF LAGUNA CHAPALA, ON MOUNTAIN SLOPE AT BORDER OF
LOWER COLORADO VALLEY, 29°27’ N, 114°25’W, 780 m, parasitic on remains of unknown
host, 6 I 1989, Wetmore 63815 (MIN); POLAND. TATRA MOUNTAINS, 1800 m, 10 X
1956, Vézda (UPS); ROMANIA. ARAD, NADLAC, NEAR THE RAILWAY STATION, 46°09’N,
20°47°E, on Acarospora nitrophila on iron-rich pebbles, 5 VII 2004, Vondrak 2134
(CBFS); SPAIN. Barcelona. MONTES MONTSENY, COLFORNIC, 1100 m, 28 V 1983,
Hladun, Poelt & Vézda (B); SWEDEN. Bohuslan. NORUM, ST. ASKERON, NEAR THE MILL,
on brown crustose lichen on roof tile, 1922, Magnusson (UPS); PAR. STENKRYA, TJORN,
OLSNAS, C. 250 M ESE OF HOLMESLATT, | m, on brown crust on silicate seashore rocks, 29
VI 2006, Westberg 06-033 (LD); Narke. PAR. GOTLUNDA, ISL. VALEN (IN LAKE
HJALMAREN), 59°15’ N, 15°45’E, 25 m, on brown crustose lichen on old roofing tile, 16
X 1993, Foucard & Nordin (UPS); Torne Lappmark. NEAR TORNE TRASK, 29 VII 1921,
Magnusson 6023 (UPS, some duplicates P simplex in other herbaria); Vastergotland.
PAR. FASSBERG, MOLNDAL, LACAREBACK, ABOVE INDUSTRIAL AREA, 93 m, on siliceous
rock outcrops, 29 VI 2006, Westberg 06-012 (LE); SWITZERLAND. Top OF NIESEN (AT
THUNNER LAKE), 2266 m, on shale rock, 1 IX 1917, Lettau (B, PARATYPE); USA.
ARIZONA: Apache Co. PETRIED FOREST NATIONAL PARK, 34°49°30”N, 109°49°00"W,
1740 m, on Candelariella rosulans, 14 V 1990, Davies 724 (ASU); CALIFORNIA: Los
Angeles Co. SANTA MONICA MOUNTAINS, TOPANGA CANYON ED EDELMAN PARK, mixed
chaparral with coastal sage scrub, 34°07°19”N, 118°35'09” W, 375 m, on sandstone, 15
VUI 2004, Knudsen 1552 w/ Sager (UCR); SANTA MONICA MOUNTAINS, LEO CARRILLO
STATE PARK, TRAIL ABOVE WILLOW CREEK, Adenostoma fasciculatum chaparral,
34°03’50°N, 118°54’56”W, 499 m, on remains probably of Acarospora oligospora, 12 XI
2005, Knudsen 4356 w/ Werth, Sagar & Heinrich (UCR); Riverside Co. MENIFEE HILLS
WILDOMAR, A. fasciculatum chaparral, 33°37'19"N, 117°12’27”"W, 590 m, on Aspicilia,
16 III 2006, Knudsen 5366 (UCR); SAN JACINTO MOUNTAINS, SAN BERNARDINO NATIONAL
FOREST, FORBES RANCH ROAD, Adenostoma sparsifolium chaparral and manzanita,
33°40'25"N, 116°36'45” W, 1651 m, on other lichens and crumbling granite, 26 HI 2006,
Knudsen 5702 (UCR); LITTLE SAN BERNARDINO MOUNTAINS, EUREKA PEAK, JOSHUA
TREE NATIONAL PARK, E AND W OF SUMMIT, pinyon pine, Quercus john-tuckeri, with
various shrubs and perennials, 34°01’57”N, 116°21°01”W, 1675 m, on rock, 22 II 2006,
Knudsen 5187 w/ Muertter, La Doux, Latman & Lee (UCR); SAN JACINTO MOUNTAINS,
BETWEEN CAHUILLA VISTA AND PINYON FLATS, juniper woodland, 33°35°08°N,
116°25°51”W, 1200 m, on lichen on granite, 10 XII 2004, Knudsen 2051 w/ Griffith
(UCR); San Bernardino Co. JOSHUA TREE NATIONAL PARK, KEYS RANCH, INDIAN COVE,
yucca, cacti, scattered scrubs and pinyon pine, 34°02’52”N, 116°10°07”W, 1260 m, on
lichen on granite, 31 II 2005, Knudsen 2602 w/ La Doux, Pietrasiak & Babich (UCR);
GRANITE MOUNTAIN, SWEENEY GRANITE MOUNTAIN RESERVE, CANYON ABOVE YUCCA
BAJADA CAMP, desert riparian and scrub, 34°48°52”N, 115°38°37”W, 1237 m, on lichen
on shale or mudstone, 12 XII 2006, Knudsen 7982 (UCR); JOSHUA TREE NATIONAL PARK,
BELLE MOUNTAIN, rocky slope with annuals, 34°00’05”N, 115°59°56”W, on granite, 22
VIUI 2005, Knudsen 3605 w/ La Doux, Galvin (UCR); SAN BERNARDINO MOUNTAINS,
SAN BERNARDINO NATIONAL FOREST, CACTUS FLATS, EAST OF SMARTS RANCH ROAD,
pinyon pine, juniper, 34°18'27"N, 116°47’27”W, 1869 m, on Acarospora strigata on
granite, 7 VI 2005, Knudsen 3327B w/ M. Knudsen (UCR); San Diego Co. ANZA
BORREGO STATE PARK, YAQUI PASS, ENCELIA, 33°08'48"N, 116°20'43”W, 550 m, on
lichens on granite, 28 IV 2006, Knudsen 5904 w/ Muertter (UCR); POINT LOMA,
CABRILLO NATIONAL MONUMENT, POINT LOMA ECOLOGICAL PRESERVE, Coastal sage
scrub, 32°40°18"N, 117°14’29”W, 116 m, on Acarospora species on sandstone, 15 V 2007,
A study of Polysporina ... 157
Knudsen 8429.1 (UCR); Santa Barbara Co. SANTA CRUZ ISLAND, CHANNEL; ISLANDS
NATIONAL PARK, ON RIDGE EAST OF CHINESE HARBOR, grassland and coastal sage scrub,
34°00°46°N, 119°35’31”W, 368 m, on thallus of Lecidea fuscoatra, 12 VI 2007, Knudsen
8523 wi I. M. & EF Brodo, Baguskus & Chaney (UCR); CONNECTICUT: Windam Co.
TOWN OF WINDHAM, W OF WINDHAM AIRPORT, SSW OF MANSFIELD HOLLOW DAM,
MANSFIELD HOLLOW STATE PARK, Ca. 100 m, Cladonia dominated gravel barren, artificial
talus slope of dam, and outwash area below dam, 28 VII 2002, Harris 46218 (NY);
COLORADO: El Paso Co. 38.7277 °N, 104.3831°W, 1725 m, locally common on Caloplaca,
4 X 2007, Morse 15885 w/ Ladd (KANU); Weld Co. NORTHWEST OF RAYMER, 91 m,
Shushan & Higgins $5647 (COLO); KANSAs: Washington Co. s END OF WASHINGTON
CO. LAKE, 39.9209°N, 97.1176°W, 427 m, 27 I 2008, Morse 16250 w/ Logan (KANU);
MICHIGAN: Washtenaw Co. OLD FIELDS NE OF CROOKED LANE, 4 VII 1976, Harris 11647
(COLO); MINNESOTA: Yellow Medicine Co. E GRANITE FALLS, UPPER SIOUX AGENCY
STATE PARK, Celtis-Quercus-Tilia forest, 44°44’02”N, 95°27'46” W, 290 m, on boulder, 5
IX 2006, Morse 14154A (UCR); NEw York: Clinton Co. POINT AU ROCHE STATE PARK,
21 V 2007, old fields, calcareous slate beach with glacial erratic, and northern hardwoods
with Thuja, initially on Acarospora, Harris 53721 (NY); PENNSYLVANIA: Bucks Co.
RALPH STOVER STATE PARK, SOUTHEAST CORNER OF PARK, BELOW STATE PARK ROAD, 82
m, extensive sandstone/ shale outcrops along creek, and shaded boulders in mixed
hemlock — hardwood forest, on thallus of Caloplaca oxfordensis on rock in stone wall, 15
IX 2007, Lendemer 9705 w/ Moroz (NY).
2. Polysporina arenacea (H. Magn.) K. Knudsen & Kocourk., comb. nov.
MycoBank MB 511742
Type: USA. UTAH: EKKER’S RANCH, 1828 m, on dry exposed, red sandstone, V 1951,
Flowers 366 (UPS, HOLOTYPE; MIN, ISOTYPE).
Basionym. Acarospora arenacea H. Magn., Acta Hort. Gétoburg. 19 (2): 33 (1952).
DESCRIPTION — THALLUS endokapylic or endolithic, vegetative hyphae hyaline,
mostly 2-3 um diam, septate, cells 2-5 um long, usually penetrating host from
below.
APOTHECIA superficial on host, brown when dry or wet, 0.7-1 mm diam,
to 1 mm tall, round or irregular, dispersed or contiguous, forming clusters of
apothecia divided by accession fissures to 3 mm wide; disc brown, convex, with
furrows, ridges, and umbos, rough and uneven, margin distinct but becoming
fissured. EXCIPULUM to 100 um thick, hyphae mostly 2-3 um diam, septate,
cells mostly 2-3 um wide, ending in expanded apices forming outer wall of
the excipulum; outer layer reddish-brown, 20-30 um thick; inner layer hyaline.
HYMENIUM 80-120 um tall, hyaline, I+ deep blue, epihymenium 10-20 um thick,
in reddish brown gel, accretions on surface of disc to 200 um high, PARAPHYSES
coherent, (1-)2-3 um diam, septate, cells 5-7 um long, shortening in upper
third, sometimes constricted at septa, infrequently branching in lower half,
unexpanded or expanded apices sometimes in darker pigment caps. Asc1 60-90
x 12-20 um, ascospores mostly 100 per ascus. AScosPoREs hyaline, simple,
(3.0-) 4.2-4.4(-7) x 2.0-3.0 um, mostly broadly ellipsoid. suBHYMENIUM
158 ... Knudsen & Kocourkova
hyaline, 30-50 um thick. HYPOTHECIUM 10-20 um thick, continuous with
vegetative hyphae. CONIDIOMATA not seen.
ECOLOGY AND SUBSTRATE — On crustose lichens on granite, sandstone, shale or
volcanic rock in open sunny habitats.
DISTRIBUTION -— Western North America (California, North Dakota, Utah)
from 40-1828 m.
Discussion - Polysporina arenacea is the only species in the genus with
distinctively brown ascomata. Otherwise, its characters overlap with the upper
range of variation in P. subfuscescens. We retain the species and transfer it from
Acarospora to Polysporina.
SPECIMENS EXAMINED — USA. CALIFORNIA: San Bernardino Co. GRANITE MOUNTAINS,
SWEENEY GRANITE MOUNTAINS RESERVE, SIBYL ALLISON TRAIL, pinyon pine and juniper
woodland with chaparral, 34°47’21"N, 115°40°18”W, 1692 m, on unknown host
on granite, 14 XII 2006, Knudsen 8055 w/ Muertter (UCR); JOSHUA TREE NATIONAL
PARK, BELOW BELLE MOUNTAINS IN UN-NAMED WASH, annuals and cacti, 34°01'08"N,
116°00°09”W, 1182 m, on unknown host on granite usually suppressing ascomata
production of the host, 22 VHI 2005, Knudsen 3620 w/ La Doux & Galvin (UCR; PRM);
Santa Barbara Co. SANTA ROSA ISLAND, CHANNEL ISLANDS NATIONAL PARK, SOUTH
SLOPE OF SOUTH PEAK, 33° 54’ 23” N 120° 7’ 42” W, 243 m, on Lecidella species and
independent on shale, 15 VIII 2007, Knudsen 8762.2 w/ Chaney & Werth (UCR); San
Luis Obispo Co. SAN SIMEON, SAN SIMEON STATE PARK, MOLINARI PROPERTY, NEXT TO
RAVINE ALONG OLD HIGHWAY ONE DIRT ROAD, on exposed slab of volcanic rock 35° 36’
14” N 121° 7 39” W, 41 m, on unknown host, 8 I 2007, Knudsen 8116 (UCR); NoRTH
DAKOTA: THEODORE ROOSEVELT NATIONAL PARK, NORTH UNIT, JUST EAST OF EDGE OF
GLACIAL OUTLOOK, 47° 37’ 04” N 103° 25’ 04” W, 670 m, on unknown white crustose
lichen, 23 VI 1998, Wetmore 80113 (MIN).
3. Polysporina pusilla (Anzi) M. Steiner ex Kantvilas, Lichenologist 30(6): 558
(1998).
Type: ITALY. IN VALLE DI FOREOLA SUPRA POSCHIAVO, ad rupem calcaream, in termino
arboreo, (Lich. Rar. Langob. Exs. No. 190) Type: not seen.
Basionym. Sarcogyne pusilla Anzi, Comment. Soc. Crittog. Ital. 1(3): 157 (1862).
= Biatorella pusilla (Anzi) Zahlbr. ex Beck, Annal. Naturhist. Hofmuseum Wein.
IV: 355 (1889).
DESCRIPTION — Lichenicolous. THALLUS endolithic or endokapylic, vegetative
hyphae hyaline, 2-3 um wide, cells 4-9 um long, I-.
APOTHECIA round, dispersed, immersed or superficial on hosts, or immersed,
emergent or sessile in rock, not apparently dividing vegetatively, mostly 0.2-0.5
(-1.0) mm diam, disc black, carbonized with umbos or ridges and furrows, -
or without carbonization and reddish when wetted with water, occasionally
white with pruina or crystals from substrate; margin progressively rugulose
with umbos, ridges and furrows. EXCIPULUM carbonized in outer layer, hyaline
to black in inner layer, 40-100 um thick, continuous with the hypothecium
A study of Polysporina ... 159
which can be 100-150 um thick, which is continuous with vegetative hyphae.
HYMENIUM (85-)100-125 um tall, hyaline, conglutinate (becoming lax in K),
PARAPHYSES mostly (1.5-)2.0 um diam, rarely paraphyses wider to 2.5-3.0 um
diam, usually mixed with thinner paraphyses, infrequent to frequent branching,
apices unexpanded or expanded to 3 tm diam, occasionally with pigment hoods
when epihymenium surface is carbonized. Epihymenium 10-25 um thick, dark
brown, in conglutinate pigment layer reddish-brown or blackish, partially
dissolving in K. ascr 65-100 x 17-25 um. ascosporEs 100-200 per ascus,
usually 4.0-5.0 x 2.0-3.0 um, ellipsoid to broadly ellipsoid. suBHYMENIUM 20-50
tum thick. HYPOTHECIUM confluent with vegetative hyphae. CONIDIOMATA not
observed.
ECOLOGY AND SUBSTRATE - In open and sunny montane habitats on calcareous
rock, apothecia immersed or superficial, parasitic on Acarospora strigata,
Caloplaca species, and Protoblastenia incrustans and probably algicolous.
DISTRIBUTION - Asia (Iran, Turkey), Europe, North America (California,
Nebraska, North Dakota), from 640 to 2800 m.
Discussion - Polysporina pusilla was originally recognized as a lichen. We
recognize it as a lichenicolous fungus with an endolithic and endokapylic
thallus. It produces apothecia immersed or superficial on a host or independent
of at least a visible host. We have a broad concept of the species recognizing
the apothecia may be as large as 1 mm diam in nutrient-rich habitats and that
apothecia are be produced in pits in limestone or on surface of rock depending
on probable substrate conditions. We do not recognize it as host specific but
it is a calciphile. It is distinguished from P subfuscescens in being a strict
calciphile and having generally thicker (2-3 vs. 1-2 um diam) paraphyses and
wider (2-3 um vs. 1.5-2.5 um) ascospores though it overlaps with higher range
of P. subfuscescens. It differs from P. arenacea in being a calciphile and having
black apothecia but is similar in paraphyses diameter and ascospore width.
As in P. subfuscescens and P. arenacea the endolithic thallus may be parasitic
on endolithic lichens or even fungi and probably on free-living algae and can
migrate through substrate to new lichen hosts. Interestingly the best-developed
specimens seen during these studies were from collections by the C. Wetmore
(MIN) and often appeared to be non-lichenicolous. In these specimens on
large-grained limestone in Nebraska and North Dakota there were abundant
free-living algae throughout upper 1 mm of the rock.
Like the lichenized Polysporina urceolata, P pusilla sometimes has a
non-carbonized disc surface in early ontogeny. Apparently in both species
carbonization of epihymenium is the continuation of the carbonization of the
excipulum and is a progressive development, usually beginning with umbos on
surface of apothecia and continuing with the formation of ridges and furrows
until it is similar to other Polysporina species.
160 ... Knudsen & Kocourkova
The species can be confused with the related lichenized species Polysporina
urceolata. They have similar spore and paraphyses sizes, but P. urceolata usually
has a lower hymenium that is 50-60 um tall (although in some specimens it
may be taller), overlapping the height of the hymenium in P. pusilla. Because
P. pusilla is a parasite, the algal layer of the host may remain for awhile, and
some specimens can only be determined through the structure of the vegetative
hyphae penetrating host from below forming a deeper structure confluent with
the hypothecium.
The species is reported new for Asia (Iran, Turkey) and North America.
SPECIMENS EXAMINED — AUSTRIA. Tirol. ALPs, 1800 m, 20 VIII 1871, Arnold (hb. J. Suza,
PRM 580306, No. 465, Anzi Exs. 190, H); 1900 m, on dolomite, 23 VII 1867, Arnold (No.
361, Anzi Exs. 190, FR); hb. A. Metzler (FR 9288); s-FACING SLOPES OF GATSCHKOPFES
ABOVE AUGOBURG MOUNTAIN COTTAGE, NW OF LANDECK, c. 2800 m, immersed in
limestone on Protoblastenia incrustans, 8 VU, 1982, Mayrhofer e& Poelt, [2] (GZU 67-82);
NORTH AND WEST OF AUGSBERG MOUNTAIN COTTAGE ABOVE GRINS, NW OF LANDECK,
2300-2400 m, 9 VII 1982, Poelt (GZU 67-82); STUBALAIER-ALPEN, GSCHNITZTAL,
TRINSER PADASTER, 2200 m, 25 VIII 1977 Schlindler e& Steiner, Plantae Graecenses Exs.
421 (B, H); IRAN. LAKE URMIAH, SHIRIN BOLAGH, 37°11'07.99"N,,. 45°23’39”E, 1290 m,
on limestone, on Caloplaca alociza, 9 V 2007, Vondrak 5142 (PRM 857464), 5438 (CBES,
UCR); USA. CALIFORNIA: Riverside Co. SAN JACINTO MOUNTAINS, SAN BERNARDINO
NATIONAL FOREST, HILLSIDE ABOVE CEDAR TRAIL, 33°39'23”N, 115°40°07”W, 1692 m,
on Caloplaca species on limestone, 29 X 2004, Knudsen 2012 (UCR); NEBRASKA: SCOTT
BLUFF NATIONAL MONUMENT, along top of SW ridge, 41°50°07”N, 103°42°14”W, 243 m,
3 VIE 1997, Wetmore 77647 (MIN); NoRTH DAKOTA: THEODORE ROOSEVELT NATIONAL
PARK, SOUTH UNIT, RIDGE NW OF BUCK HILL ON ROCK LEDGES AND IN PRAIRIE, 847 m,
no lichen host apparent, 26 VII 1982, Wetmore 45159 (MIN); TURKEY. Bitlis. Lake
VAN, 4.5 KM SW OF ADILCEVAZ, 38°47°00"N, 42°41°00.68”E, on Caloplaca erodens & C.
albopruinosa with Opegrapha vulpina, 12 V 2007, Vondrak 5139 (PRM 857465).
Species excluded from the genus Polysporina
4. Sarcogyne lapponica (Ach. ex Schaer.) K. Knudsen & Kocourk., comb. nov.
MycoBank MB 511743
Type: EUROPE. on wood, H-ACH 178 (LECTOTYPE designated here).
Basionym. Lecidea lapponica Ach. ex Schaer., Lichen. Helvet. Spicil., p. 205 (1833).
= Biatorella lapponica (Ach. ex Schaer.) Almq., Botan. Notiser, p. 67 (1866).
= Acarospora lapponica (Ach. ex Schaer.) ‘Th. Fr., Lich. Scand., p. 218 (1871).
= Myriospora lapponica (Ach. ex Schaer.) Hue, Morph. et Anat., p. 165 (1909).
= Polysporina lapponica (Ach. ex Schaer.) Degel., Acta Reg. Soc. Sci. Litt. Gothob.,
Bot. ser. 2: 103 (1983 [‘1982’]).
= Acarospora tromsoeensis Norman, Kgl. Norske Vidensk.-Selsk. Skritter, Vol. 5., p. 340
(1868). Type: NORWAY. Troms. TROMs@, Norman (W, LECTOTYPE designated
here; S, UPS, ISsOLECTOTYPES).
DESCRIPTION — Non-lichenicolous, lichenized. THALLUS endolithic in rock
or thin ochraceous crust in surface of wood, 20-30 um thick, composed of
thin intricate hyphae, indeterminate and discontinuous, sometimes reduced to
small area around or below apothecium.
A study of Polysporina ... 161
APOTHECIA black, 100-400(-500) um diam, 150-300 um tall, in fertile
globose or subglobose stromata opening more or less to expose apothecial
disc, sessile, dispersed, rarely contiguous, with no evidence of vegetative
division. Disc pale-reddish to yellow-brown, flat, without carbonization of the
epihymenium, 40-300 tm diam, sometimes best seen when wetted. Thalline
margin lacking, margin carbonized, smooth to rough, more or less fissured,
but lacking distinct joint lines as in S. privigna. EXCIPULUM derived from
ontogeny of the wall of globose infertile stroma, up to 100 um thick, outer layer
ca. 20-50 um thick, of indistinct carbonized hyphae, the inner layer 20-50 um
thick, containing crystals, distinct or indistinct from an innermost layer of thin
prosoplectenchyma continuous with hypothecium. Indistinct medullary area
beneath hypothecium mixed with substrate. Photobiont, chlorococcoid green
alga, 6-16 um diam, not forming a clear stratum in thallus or under apothecia,
sometimes not evident. HYMENIUM 100-140 um tall, hyaline, epihymenium
dark, conglutinate, 20 um thick. PARAPHYSES 1 «tm diam, coherent, with some
branching, apices unexpanded. ascr clavate 50-75 x 15-22 um, Acarospora-
type. ASCOSPORES about 100 per asci, simple, hyaline, narrowly ellipsoid, 2.5-5 x
1-1.5 um. SUBHYMENIUM 20-35 um thick, hyaline, subparaplectenchyamatous.
HYPOTHECIUM is 10-15 um thick, somewhat obscure beneath subhymenium.
CONIDIOMATA not observed, but probably in young infertile globose stromata.
ECOLOGY AND SUBSTRATE — In open habitat on slate or gneiss, probably rarely
on wood, non-lichenicolous, sometimes occurring with Polysporina simplex.
DISTRIBUTION — Europe (Norway, Sweden, Switzerland).
Discussion — The systematic and nomenclatural history of Sarcogyne lapponica
has seen it placed in five genera. Schaerer (1833) originally described the
species as a Lecidea based on concepts in his period. Almquist (1866) placed it
in Biatorella. Fries (1871) transferred it to Acarospora. The description of Hue
(1909) is based on the Acharius types and he transferred the species to the
genus Myriospora, which is currently monotypic, containing only M. heppii
(Nageli ex Korb.) Hue (Harris & Knudsen 2006, Knudsen 2008c). Degelius
(1986) transferred the name to Polysporina and applied it to the lichenicolous
taxon discussed in this paper as P. subfuscescens. Examination of the type for
this paper resulted in the discovery that the Acharius specimen is lichenized
and lignicolous and that P lapponica is actually a Sarcogyne species that is
totally unrelated to the lichenicolous taxon we recognize in this treatment as
P. subfuscescens.
The development of the ascocarp in S. lapponica occurs in two stages. In the
first stage an infertile globose stroma develops, probably containing pycnidia.
In the second stage, the stroma becomes fertile, more or less opening into a
flat apothecium. The two-stage development is characteristic of a number of
species of the core group of Sarcogyne (S. clavus, S. privigna, and S. reebiae
162 ... Knudsen & Kocourkova
K. Knudsen). Sarcogyne s. str. differs from Polysporina primarily in not
developing carbonization of apothecial surface. Because Lecidea lapponica
has both the two-stage development of the ascocarp and lacks epihymenial
carbonization, it is transferred to the genus Sarcogyne.
Magnusson’s concept of Sarcogyne lapponica as Acarospora lapponica
(Magnusson 1924, 1929) includes the heterogeneous Lecanora belonioides Nyl.,
which he synonymized with A. lapponica. Lecanora belonioides was described
from a specimen from Japan and is similar to taxa of Nylander’s broad concept
of Acarospora fuscata (Schrad.) Arnold (Nylander 1864), and the combination
Acarospora belonioides (Nyl.) Arnold was made. In the Asian specimens of
L. belonioides we examined (H!, S!), there were at least two taxa, one a definite
Acarospora and the other a probable Sarcogyne. We reject the synonymization of
L. belonioides with S. lapponica (Magnusson 1924, 1929). Lecanora belonioides
needs to be revised beginning with fresh and ample collections from Japan, the
type area. Magnusson’ reports of S. lapponica occurring in Asia are based on
L. belonioides. The distribution of S. lapponica is aOR) recognized as being
restricted to Norway, Sweden, and Switzerland.
Making ample field collections on rock is problematic because not only
is Sarcogyne lapponica minute but it also sometimes occurs with Polysporina
simplex. When the ascocarps of S. lapponica are not fully open exposing the
disc it would be hard to distinguish from P. simplex, especially when the
excipulum becomes rough and somewhat fissured. S. lapponica is distinguished
from P. simplex by the lack of epihymenial carbonization and usually globose
ascocarps, otherwise the internal structure of the hymenium and the size of
ascospores are the same. In some of specimens we examined Magnusson mis-
determined P simplex as S. lapponica.
In the Swedish key for Sarcogyne (Foucard 2001) the couplet distinguishing
S. clavus and S. privigna would need to be revised adding and distinguishing
S. lapponica as having apothecia 0.1-0.4 mm, grouped with S. privigna, and
having a hyaline hypothecium.
Another problem in identifying Sarcogyne lapponica is that S. algoviae
H. Magn. has been reported from Fennoscandia (Santesson et. al. 2004).
Sarcogyne algoviae is a member of the core group of Sarcogyne and is
distinguished by the small size of its apothecia (0.3-0.5 um diam) and its
hyaline hypothecium. It has overlapping measurements of the paraphyses and
ascospores with S. lapponica and P. simplex, but while both those species have
hymenia generally above 100 um tall, S. algoviae has a hymenium 50-60 um
tall. Based on the specimens of S. algoviae that Claude Roux supplied to us, it is
distinguished from S. lapponica by a highly carbonized and fissured excipulum
with somewhat flattened ascocarps immersed in the substrate and in being
a calciphile usually occurring on dolomite. Sarcogyne algoviae needs further
study.
A study of Polysporina ... 163
SPECIMENS EXAMINED ~ SWITZERLAND. 12 VII 1921, Frey (UPS); SWEDEN.
Vastergétland. FLosy, VIII 1915, Vrang [2] (UPS); Torne Lapmark. par. JUKKASJARVI,
BJORKLIDEN, VII 1927, Vrang (S); ABISKO DISTRICT, on slate, 2 VIII 1921, Magnusson
6165 (S); 1924, Malme (S).
Conclusion
Future revisions of the lichenicolous Polysporina would benefit from
molecular studies at the genus and family level. A robust molecular phylogeny
could corroborate the species concepts defined in this paper, determine the
placement of Polysporina within the family Acarosporaceae and its relationship
to Sarcogyne s. str., and give insight into the loss of lichenization within the
family Acarosporaceae.
In Acarosporaceae, narrow species concepts do not hold up well when
taxonomic studies are done of a larger set of multi-regional specimens. Future
description of new taxa in Polysporina should not be based on Magnusson’s
narrow species concepts.
Acknowledgements
Thanks to Anders Nordin (UPS) and James C. Lendemer (NY) for reviewing the
manuscript. We thank the curators of the herbaria who supplied us with specimens.
We thank Claude Roux and Valérie Reeb for their valuable comments on drafts of this
manuscript.
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Vézda A. 1970. Neue und wenig bekannte Flechten in der Tschechoslowakei. I. Geobotanica et
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MYCOTAXON
Volume 105, pp. 165-166 July-September 2008
Nomenclatural note on an Asian Pluteus
ANDREW M. MINNIS’ & WALTER J. SUNDBERG?
‘Drew. Minnis@ars.usda.gov
Systematic Mycology & Microbiology Laboratory
USDA-ARS, Rm. 304, BO11A, 10300 Baltimore Avenue
Beltsville, MD 20705, USA
* sundberg. wj.407@verizon.net
107 Cardinal Drive, Murphysboro, IL 62966-5255, USA
Abstract — Asian Pluteus taxa are not as well known as those from Europe and North
America. Pluteus australis Z.S. Bi, one of the taxa described as new from Asia, is an
illegitimate later homonym of Pluteus australis Murrill, a species described as new from
the USA. Hence, the new name, Pluteus bii, is presented for the former species.
Key words — Agaricales, Basidiomycota, nomenclature, Pluteaceae
Introduction
Pluteus Fr., an agaric genus, is a circumglobal saprotroph and commonly
lignicolous (Singer 1986). Relatively little is known about the Pluteus mycota
of China and Korea. For such a large area with a diverse flora, few new species
of Pluteus have been described. In fact, based on available literature, Pluteus
luteus (Redhead & B. Liu) Redhead (Redhead 1984) and Pluteus australis
Z.S. Bi (Bi 1988), both from China, are the only Pluteus taxa reported as new
from this area. Lee et al. (1992) reported on the Korean Pluteus mycota, but
they did not recognize any new taxa and none of the species they listed were
described as new from Asia.
Bi (1988) was unaware when he discovered and named Pluteus australis
Z.S. Bi, a species in Pluteus section Hispidoderma Fayod, that Pluteus australis
Murrill, a species in Pluteus section Pluteus (Banerjee & Sundberg 1995),
had already been described from the USA (Murrill 1945). According to
the International Code of Botanical Nomenclature (McNeill et al. 2006),
Pluteus australis Z.S. Bi is an illegitimate later homonym. Because of this, a
replacement name is needed and, the following new name is proposed in honor
of Bi Zhi-Shu.
166 ... Minnis & Sundberg
Taxonomy
Pluteus bii Minnis & Sundb., nom. nov.
MycoBAnk 511726
= Pluteus australis Z.S. Bi, Acta Mycol. Sin. 7: 90, 1988, non Pluteus australis Murrill,
Proc. Florida Acad. Sci. 7: 119, 1945 (“1944”).
Acknowledgments
We thank Drs. Else C. Vellinga and Olivia Rodriguez Alcantar of the University of
California at Berkeley and Universidad de Guadalajara, respectively, for their reviews
of this manuscript.
Literature cited
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Bi Z-S. 1988. One new species of Pluteus Fr. from Guangdong Province of China. Acta Mycol. Sin.
1: 89-92.
Lee J-N, Lee H-K, Min K-H, Park W-H, Kim Y-S. 1992. Studies on genus Pluteus of Korea. Korean
J. Mycol. 20: 296-301.
McNeill J, Barrie FR, Burdet HM, Demoulin V, Hawksworth DL, Marhold K, Nicolson DH, Prado
J, Silva PC, Skog JE, Wiersema JH, Turland NJ (eds.). 2006. International Code of Botanical
Nomenclature (Vienna Code): Adopted by the Seventeenth International Botanical Congress,
Vienna, Austria, July 2005. ARG Gantner, Ruggell. 568 p.
Murrill WA. 1945 (“1944”). New Florida fungi. Proc. Florida Acad. Sci. 7: 107-127.
Redhead SA. 1984. Mycological observations, 4-12: on Kuehneromyces, Stropharia, Marasmius,
Mycena, Geopetalum, Omphalopsis, Phaeomarasmius, Naucoria and Prunulus. Sydowia
37: 246-270.
Singer R. 1986. The Agaricales in Modern Taxonomy. 4" ed. Koeltz Scientific Books, Koenigstein.
981 p.
MYCOTAXON
Volume 105, pp. 167-170 July-September 2008
The macrofungal checklist of Liguria (Italy):
current survey status
MircA ZOTTI’*, ALFREDO VIZZINI 7, MIDO TRAVERSO?,
FABRIZIO BOCCARDO*, MARIO PAVARINO'! & MAURO GIORGIO MARIOTTI!
*milla@klaatu.com.dist.unige. it
'DIP.TE.RIS - Universita di Genova - Polo Botanico “Hanbury”
Corso Dogali 1/M, 116136 Genova, Italy
? MUT- Universita di Torino, Dipartimento di Biologia Vegetale
Viale Mattioli 25, 110125 Torino, Italy
°Via San Marino 111/16, 116127 Genova, Italy
‘Via E. Bettini 14/11, 116162 Genova, Italy
Abstract — The goal of this paper is to integrate and update the first edition of the
checklist of Ligurian macrofungi with data resulting from mycological research
conducted mainly in holm-oak woods during the past last three years. Of the 172
new taxa collected, 15 represent Ascomycota, 157 represent Basidiomycota; 12 taxa are
recorded for the first time from Italy and many others are considered rare or infrequent.
Each taxonomic entry includes Latin name, author, habitat, height, and WGS-84 Global
Position System (GPS) coordinates. This work, together with the original Ligurian
checklist, represents a contribution to the national checklist. The complete checklist is
available at http://www.mycotaxon.com/resources/weblist.html.
Key words — mycological flora, new reports, species richness
Liguria represents a very interesting region from a mycological point of
view. Zotti & Orsino (2001) have noted that its macrofungi, directly and
indirectly associated with the vegetation, are frequent, abundant, and highly
diverse. Observations support a high level of fungal biodiversity (sometimes
called “mycodiversity”) for Liguria, which covers only about 2% of the Italian
territory but with 36% of all species recorded in Italy represented. Moreover,
species richness is likely underestimated, as only 50% of Liguria was actually
investigated (see Fig. 1 in the web Checklist). It is significant that such a brief
study carried out in such a limited number of new areas increases so greatly the
total number of species observed. The new data used to update the previous
Ligurian checklist (Zotti & Orsino 2001) and Italian checklist (Onofri et al.
2005) are based primarily on collections made in previously unsurveyed holm-
168 ... Zotti & al.
oak woods (Quercus ilex L.). Statistical analyses of some sites by Zotti (2004)
confirm the high level of mycodiversity of the region. The paper reports data
of mycological investigations carried out through the last six years (2001-07).
Moreover, a brief annex is also included that lists the species recorded in Liguria
and described in recent papers.
MeETHODOLOGY—Each collection was identified based on macro- and
micromorphological characters. References consulted included Alessio &
Rebaudengo (1980), Antonini & Antonini (2002), Bas et al. (1988, 1990, 1995,
1999), Basso (1999), Bernicchia (2005), Bon (1984), Breitenbach & Kranzlin
(1984), Consiglio (1996), Consiglio et al. (2003, 2004, 2005), Contu (2000,
2003), Eyssartier & Buyck (2000), Fontenla et al. (2003), Fraiture (1993), Froslev
et al. (2006), Grupo Ibero-insular de Cortinariologos (GIC 2007), Jiilich (1989),
Kreisel (1987), Ladurner & Simonini (2003), Malencon & Bertault (1970),
Moser (1980), Noordeloos et al., (2001), Raillére & Gannaz (1999), Riva (1988,
2003), Robich (1990), Romagnesi (1967), Sarnari (1998, 2005), Stangl (1991),
Traverso (1999), Vesterholt (2002). A Garmin (eTrex Summit) Global Position
System (GPS) was used to determine collection sites in WGS-84 coordinates in
decimal degrees. Family, order and class names follow Kirk et al. (2001), Vizzini
(2004), and Hibbett (2006). Abbreviation of author names follows Brummitt &
Powell (1992) and IPNI (2008). Several species cited in the previous checklist
have since been documented by Balletto (1972). The new records are intended
to add previously omitted collection and site information (indicated with an
asterisk in the taxonomic list). Species flagged with “KM” represent taxa not yet
cited in Italian checklist (Onofri et al. 2005). Finally, the annex reports a group
of species recently observed and the related bibliographic references.
RESULTS—172 taxa, representing 58 genera and 35 families, were recorded. The
list is arranged alphabetically in order to simplify the seeking and the reading
of species. Each taxonomic entry provides the Latin name, authority, WGS-84
Global Position System (GPS) coordinates in decimal degrees, altitude, survey
date when the species was first found, and collection site habitat.
ConcLusions— The weblist lists new species observed in Liguria (primarily
from holm-oak habitats) during the 2001-07 mycological surveys. The
newly recorded 172 taxa represent Basidiomycota and Ascomycota. The paper
emphasizes the preliminary nature of mycological research in Liguria and
draws attention to its unusually high level of fungal richness. Although the areas
where the species were collected represent a negligible portion of the whole °
Ligurian territory, the increase of observed species comprises approximately
13% of all the records of the previous list. Furthermore, 12 basidiomycetes —
Cortinarius chevassutii Rob. Henry, C. ionochlorus var. leucophyllus Malencon,
C. murellensis Cors. Gutiérrez et al., C. parasuaveolens (Bon & Trescol) Bidaud
Macrofungi of Liguria (Italy) ... 169
et al., C. sancti-felicis Froslev & T.S. Jeppesen, C. veraprilis Chevassut et al.,
Inocybe squarrosa Rea, Laccaria macrocystidiata (Migl. & Lavorato) Pazmany,
Melanoleuca amica (Fr.) Singer, M. grammopodia var. obscura Bon, Russula
subazurea Bon, Xerula radicata var. alba Dorfelt — have not been previously
cited in the Italian checklist (Onofri et al. 2005). These facts should encourage
studying new territories and habitats. Finally, taxonomic reevaluations of
complex genera such as Russula and Cortinarius might uncover species not
previously recognized in the country.
Acknowledgments
The authors like to thank the reviewers Marco Contu and Gabriel Moreno (Dpt. Biologia
Vegetal Univ. Alcala de Henares) for their helpful and constructive comments.
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Candusso M. 1997. Fungi Europaei 6. Hygrophorus s.1. Libreria Basso, Alassio.
Consiglio G. 1996. Cortinarius catharinae sp. nov. RAM 39(3): 195-200.
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Contu M. 2000. Saggio di una chiave per la determinazione delle specie del genere Amanita
osservate in Sardegna. BGMB 43(2): 67-6.
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46(1): 5-8.
Eyssartier G, Buyck-B. 2000. Le Genre Cantharellus en Europe. Nomenclature et Taxonomies. Bull.
Soc. mycol. Fr. 116(2): 91-137.
Fontenla R, Gottardi M, Para R. 2003. Osservazioni sul genere Melanoleuca. Fungi non delineati.
Ed. Candusso. Alassio
Fraiture A. 1993. Le Amanitopsis d Europe. Opera Bot. Belg. 5, Meise.
Froslev TG, Jeppesen TS, Lzessge T. 2006. Seven new calochroid and fulvoid species of Cortinarius.
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Grupo Ibero-insular de Cortinariologos (GIC 2007). Cortinarius Ibero-insulares-1. Fungi Non
Delineati, Pars XLI-XLII, Ed. Candusso, Alassio, Italy.
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Onofri S, Bernicchia A, Filipello V, Padovan F, Perini C, Ripa C, Salerni E, Savino E, Venturella G,
Vizzini A, Zotti M, Zucconi L. 2005. Checklist of Italian fungi. Carlo Delfino Editore, Sassari.
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Romagnesi H. 1967. Les Russules d’Europe et d'Afrique du Nord. Editions Bordas, Paris.
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Traverso M. 1999. Il genere Amanita in Italia. A.M.E.R., Roma.
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MY COTAXON
Volume 105, pp. 171-174 July-September 2008
Gloeoporus dichrous var. niger comb. nov.
MICHAL TOMSOVSKY! & LEIF RYVARDEN?
tomsovsk@mendelu.cz
‘Faculty of Forestry and Wood Technology
Mendel University of Agriculture and Forestry in Brno
Zemédeélskd 3, CZ-613 00, Brno, Czech Republic
*Botanical Institute, University of Oslo
PO. Box 1066, Blindern, N-0316 Oslo, Norway
Abstract — Molecular phylogenetic study of Ceriporiopsis nigra, a species known
only from its type locality, was performed to reveal its evolutionary relationship to
Ceriporiopsis pannocincta and to the type species of the genus, C. gilvescens. The
sequencing of ITS and mitochondrial SSU regions of ribosomal DNA revealed identity
of the sequences of C. nigra and Gloeoporus dichrous, apart from Ceriporiopsis. Even
though C. nigra and G. dichrous share similar microscopy, the two species differ
remarkably in colour of basidiocarp. Thus, the new combination Gloeoporus dichrous
var. niger is proposed.
Key words — Basidiomycota, taxonomy, polyporoid clade
Introduction
Although the European mycobiota is relatively well investigated, every now
and then new species are still described at the beginning of the 21th century.
An example of such a species is Ceriporiopsis nigra (Ryvarden 2003), to date
known only from the type locality. The species was placed in Ceriporiopsis due
to its resupinate, poroid hymenophore with monomitic hyphal system and
clamps. C. nigra seems to be related to Ceriporiopsis pannocincta (Gloeoporus
pannocinctus). Both species share a distinct resinous layer between the tubes
and context and similar shape of basidiospores, but they differ in pore width.
Due to a probable polyphyletic origin of Ceriporiopsis as currently conceived
(Ryvarden & Gilbertson 1993), we decided to test the taxonomic position of
the included species by molecular methods comparing C. nigra not only with
C. pannocincta but also with the generic type species C. gilvescens.
Material and methods
To assure the taxonomic position of Ceriporiopsis nigra sequencing of the ITS region
of nuclear ribosomal DNA (ITS) and small subunit of mitochondrial ribosomal DNA
(mitSSU) were carried out.
172 ...TomSovsky & Ryvarden
The holotype of C. nigra (deposited in O herbarium; Ryvarden no. 17528) was sequenced
together with collections of Ceriporiopsis gilvescens and C. pannocincta (collected in the
Czech Republic, deposited in BRNM herbarium; specimens no. 709970 and 709972,
respectively).
DNA was isolated from the dried tissue of herbarium specimens using PowerSoil™
DNA Isolation Kit (Mo-Bio). For amplification of ITS and mitSSU the primer pairs
ITS1/ITS4 and MS1/MS2 were used (White et al. 1990). The DNA was amplified with
PCR, using the Mastercycler® ep thermocycler (Eppendorf). The PCR amplifications
were performed according to TomSovsky et al. (2006). The only modification was lower
temperature of mitSSU region amplification (50 °C) instead of 55 °C used for ITS one.
The PCR products were purified by NucleoSpin Extract II (Macherey-Nagel) prior to
sequencing.
Sequences were determined with an ABI PRISM 3100 Avant DNA sequencer
(Applied Biosystems) at the Department of Animal Morphology, Physiology and
Genetics, Faculty of Agriculture, Mendel University, Brno using the ABI PRISM BigDye
terminator v1.1 cycle sequencing kit (Applied Biosystems). All samples were sequenced
with the primers used in the PCR.
GenBank search of newly obtained mitSSU sequence of C. nigra using BLAST
(Altschul et al. 1990) resulted in 100% identity with those of Gloeoporus dichrous
(GenBank Access. No. AY986495). Due to this result, the collection of G. dichrous
(collected in Slovakia, deposited in BRNM herbarium; specimen no. 709971) was also
sequenced to assure the phylogenetic proximity of the two species. The sequences were
deposited in the EMBL Nucleotide Sequence Database, their GenBank ITS and mitSSU
accession numbers are as follows: Ceriporiopsis nigra (EU546098, EU546101), C.
gilvescens (EU546104, EU546103), C. pannocincta (EU546099, EU546102), Gloeoporus
dichrous (EU546097, EU546100).
All sequences were edited manually using BioEdit version 4.7.1. (Hall 1999) and
two sequences from the GenBank (Gloeoporus dichrous mitSSU sequence, AY986495;
Ceriporiopsis pannocincta ITS one, AY219361) were added to the dataset. The combined
ITS and mitSSU alignment with introduced gaps consisted of 1088 characters including
830 constant, 258 variable and 68 parsimony informative sites. The length of mitSSU
sequence of C. gilvescens differs apparently from the other studied Ceriporiopsis species
(1525 bp instead of 452-455 bp) due to presence of two introns, and so these heterologous
regions were excluded from the study prior to aligning.
Bayesian analysis was performed using software MrBayes 3.0 (Huelsenbeck &
Ronquist 2001) while the maximum parsimony analysis (MP) was performed with
PAUP 4.0 (Swofford 1999). The support for the topology of MP trees was estimated
using 1000 bootstrap-replicates. In MrBayes, 2 x 10° generations of the Markov Chain
Monte Carlo were run with four simultaneous chains, heating temperature 0.2. First
2000 trees were discarded as burn-in.
Results and discussion
The newly obtained Ceriporiopsis nigra sequences of both gene regions show
100% identity to those of Gloeoporus dichrous. The phylogenetic analysis of
combined dataset resulted in the same phylogram topologies with respect to
Gloeoporus dichrous var. niger comb. nov. ... 173
Ceriporiopsis gilvescens
EU546104/ EU546103
Ceriporiopsis pannocincta
400/1.00 AY219361/ -
Ceriporiopsis pannocincta
EU546099/ EU546102
Gloeoporus dichrous
-| AY9864395
100/1.00
Gloeoporus dichrous
EU546097/ EU546100
50/0.50
Ceriporiopsis nigra
EU546098/ EU546101
0.1
Figure 1. Phylogenetic tree of Ceriporiopsis nigra and related taxa based on combined sequences of
the ITS and mitSSU regions of nuclear ribosomal DNA region. The unrooted tree was constructed
by the Bayesian analysis. Bootstrap values from Maximum Parsimony and Bayesian posterior
probabilities, respectively, are shown at the nodes. ‘The bar indicates number of substitutions per
position.
the used MP and Bayesian analyses (Fig. 1). The MP analysis gave 278 most
parsimonious trees; tree length = 278, consistency index (CI) = 1.000, and
retention index (RI) = 1.000.
Analyses revealed that C. nigra is more closely related to Gloeoporus dichrous
than to the type species of Ceriporiopsis. Ceriporiopsis pannocincta is also more
proximal to the C. nigra / Gloeoporus dichrous clade than to C. gilvescens.
Accordingly a new combination is proposed:
Gloeoporus dichrous var. niger (Ryvarden) TomSovsky & Ryvarden, comb. nov.
MycoBank : MB 511831
Basionym: Ceriporiopsis nigra Ryvarden, Windahlia 24: 26, 2001, as ‘niger.
Basidiocarps annual, resupinate, brittle when dry, 1-2 mm thick, margin narrow
and white, pore surface first ochraceous becoming blackish brown, pores thin-
walled, angular, 2-3(—4) per mm, tube layer concolorous, resinous and brittle
up to 1 mm deep, black resinous zone present above the tubes, about 0.1 mm
thick, subiculum white, up to 0.5 mm thick.
174 ...TomSovsky & Ryvarden
Hyphal system monomitic, generative hyphae hyaline, thin to slightly thick-
walled, with clamps and branched, negative in Melzer’s reagent, 2-6 um in
diam. Cystidia and other sterile hymenial elements absent. Basidia clavate, four-
sterigmate, 12-14 x 3.5-4.5 tum, with a basal clamp. Basidiospores allantoid,
hyaline, smooth, negative in Melzer’s reagent, 3.7-4.0 x 0.8-1.0 um. Causing
white rot of dead lying trunk of Populus tremula. Known only from the type
locality in Ostfold county, Norway.
Gloeoporus dichrous var. niger and G. dichrous have identical morphological
and molecular characters, e.g. basidiocarp morphology, hyphal system and
basidiospore dimensions (Ryvarden & Gilbertson 1993, Ryvarden 2001). On
the other hand, Gloeoporus niger lacks the apparent reddish to purplish tint
of pore surface so typical for G. dichrous but this character is not sufficient
to discriminate the fungi at the specific level. The occurrence of intermediate
forms between the new described variety and G. dichrous s.s. has not yet
been verified. If populations concerning both taxa would be discovered, the
interfertility between them should be tested to reveal a possible intersterility.
Acknowledgments
The work was supported by the Czech Science Foundation, project no. 526/06/P017. We
are grateful to Dr. Annarosa Bernicchia (University of Bologna, Italy) and Dr. Tuomo
Niemela (Finnish Museum of Natural History, University of Helsinki, Finland) for
presubmittal review of the manuscript.
Literature cited
Altschul SF, Gish W, Miller W, Myers EW, Lipman DJ. 1990. Basic local alignment search tool.
J. Mol. Biol. 215:403-410.
Hall TA. 1999. BioEdit: a user-friendly biological sequence alignment editor and analysis program
for Windows 95/98/NT. Nucleic Acids Symp. Ser. 41: 95-98.
Ronquist EF, Huelsenbeck JP. 2003. Mr. Bayes3: Bayesian phylogenetic inference under mixed
models. Bioinformatics 19: 1572-1574.
Ryvarden L, Gilbertson RL. 1993. European polypores, Part 1. Fungiflora: Oslo (Norway): 393 pp.
Ryvarden L. 2001. Ceriporiopsis niger nova sp. Windahlia 24: 25-27.
Swofford DL. 1999. PAUP* Phylogenetic analysis using parsimony. Version 4. Sinauer:
Sunderland (USA).
Tomsovsky M, Kolarik M, Pazoutova S, Homolka L. 2006. Molecular phylogeny of European
Trametes (Basidiomycetes, Polyporales) species based on LSU and ITS (nrDNA) sequences. Nova
Hedwigia 82: 269-280. ;
White TJ, Bruns TD, Lee SB, Taylor JW. 1990. Amplification and direct sequencing of fungal
ribosomal RNA genes for phylogenetics, pp. 315-322. In: PCR Protocols, A guide to methods
and applications (Innis MA, Gelfand DH, Sninsky JJ, White TJ, eds.). Academic Press: New
York (USA).
MYCOTAXON
Volume 105, pp. 175-184 July-September 2008
Notes on Heteroconium and a new species from Venezuela
RAFAEL F, CASTANEDA RUIZ
rfcastaneda@inifat.co.cu
Instituto de Investigaciones Fundamentales en Agricultura Tropical Alejandro de Humboldt
Calle 1 Esq. 2, Santiago de Las Vegas, C. Habana, Cuba, C.P. 17200
TERESA ITURRIAGA
titurri@usb.ve
Departamento de Biologia de Organismos, Universidad Simon Bolivar,
Apartado 8900. Sartanejas, Baruta, Edo Miranda, Venezuela
GABRIELA HEREDIA ABARCA
gabriela. heredia@inecol.edu.mx
Instituto de Ecologia, A.P. 63, Km 2.5,
Antigua carretera a Coatepec, Xalapa, Veracruz, México
Davip W. MINTER
d.minter@cabi.org
CABI, Bakeham Lane, Egham, Surrey, TW20 9TY, United Kingdom
JOSEPA GENE
josepa.gene@urv.cat
Unitat de Microbiologia, Facultat de Medicina i Ciencies de la Salut,
Universitat Rovira i Virgili 43201 Reus, Spain
MARC STADLER
marc.stadler@t-online.de
University of Bayreuth, Faculty of Biology, Chemistry and Earth Sciences,
Dept. Mycology, D-95440 Bayreuth, Germany
MASATOSHI SAIKAWA
saikawa @u-gakugei.ac.jp
Department of Biology, Tokyo Gakugei University
Nukuikita-machi, Koganei-shi, Tokyo 184-8501, Japan
CAROLINA SILVERA-SIMON
mariacarolina.silvera@urv.cat
Unitat de Microbiologia, Facultat de Medicina i Ciencies de la Salut,
Universitat Rovira i Virgili 43201 Reus, Spain
Abstract — Heteroconium avilae sp. nov., found on the decaying rachis of an
unidentified palm tree in “Cerro El Avila” near Caracas, Venezuela, is described and
illustrated. Heteroconium avilae is characterized by cylindrical to fusiform, sometimes
176 ... Castaheda Ruiz & al.
slightly navicular, 1- to 3- or rarely 4-septate, smooth conidia that have dark brown
to brown central cells and pale brown end cells. Illustrations of conidia of published
Heteroconium species are provided. A key to Heteroconium species is presented.
Key words — anamorphic fungi, systematics, tropical rainforest
Introduction
During three expeditions in 2006 through Venezuela’s “Cerro El Avila”
rainforest, 102 anamorphic fungi were examined from plant material on soil or
submerged in water, and a conspicuous fungus from the genus Heteroconium
was collected. The specimens showed differences from previously described
species and are therefore described as a new species.
Materials and methods
Samples of plant litter collected in separate paper bags were taken to the
laboratory where each was incubated at 25° C in Petri dishes within 50 L
plastic moist chambers containing 200 ml sterile water and 2 ml glycerol. The
plant material was screened at regular intervals for microfungi. Mounts were
prepared in polyvinyl alcohol-glycerol (8 g per 100 ml H,0, plus 5 ml glycerol),
and measurements were made at a magnification of x 1000.
Taxonomy
Heteroconium avilae R.F. Castafieda, Iturr., Heredia & Minter anam. sp. nov.
MycoBank MB 511192 Fics. 1-10
CoLonliaE in substrato naturali effusae, nigrae. Mycelium partim superficiale et partim
in substrato immersum, ex hyphis septatis, laevibus, atrobrunneis vel brunneis, 1-2
yum diam., compositum. CONIDIOPHORA macronematosa, mononemata, erecta, recta,
cylindrica, 6- usque ad 14-septata, laevia, 180-375 x 16-25 um, nigra vel atrobrunnea ad
basim et pallide brunnea ad apicem. CELLULAE CONIDIOGENAE hologenosae, uniloculosae,
laeves, dilute brunneae, 18-25 x 6-8 um, in conidiophoris incorporatae, interdum cum
1-2 proliferationibus percurrentibus cylindricis praeditae. SECEDENTIA CONIDIORUM
schizolytica. Conrp1A cylindrica usque fusiformia, interdum leviter navicularia, 1- ad
3- (rarissime 4-) septata, utrimque truncata 23-33 x 8-12 um, cellulis cum centralibus
brunneis et utrimque dilute brunnea, laevia, sicca, blastocatenulata cum catenis fasciaria
formata. TELEOMORPHOSIS: ignota.
TYPUS: Venezuela, Caracas, “Cerro El Avila’, on decaying rachis of an unidentified
palm tree. Coll. A. Fernandez, 17.V1.2006. Holotype: INIFAT C06/47-1. Isotype: USB
C06/47-1.
Erymo.oey: Latin, avilae, in reference to “El Avila? mountain from Caracas,
Venezuela.
COLONIES on the natural substratum effuse, black. Mycelium superficial
and immersed, composed of septate, smooth-walled, dark brown or brown
hyphae, 1-2 um diam. CONIDIOPHORES macronematous, mononematous,
Heteroconium (Venezuela) ... 177
Figs. 1-10 Heteroconium avilae, from holotype (INIFAT C06/47-1). Figs. 1-4, 10. Conidia. Figs.
5-9. Conidiogenous cells and conidial chains. Scale is indicated by bars = 10 um.
178 ... Castafeda Ruiz & al.
erect, straight, unbranched or with a secondary branch near the proliferation,
cylindrical, 6- to 14-septate, smooth-walled, 180-375 x 16-25 um, black to dark
brown at the base and pale brown towards the apex. CONIDIOGENOUS CELLS
hologenous, unilocal, smooth-walled, pale brown, 18-25 x 6-8 tum, integrated,
with 1-2 cylindrical percurrent proliferations. CONIDIAL SECESSION schizolytic.
ConipiA cylindrical to fusiform, sometimes slightly navicular, truncate at the
base and apex, but the distal conidium rounded at the apex, 1- to 3-septate,
rarely 4-septate, 23-33 x 8-12 um, the central cells brown to dark brown, the
end cells very pale brown, guttulate, smooth-walled, dry, blastocatenulate with
band-shaped pigmentation. TELEOMORPH: unknown.
CoMMENTs: The genus Heteroconium was erected by Petrak (1949) for the
type species, H. citharexyli Petr., and is distinguished by macronematous,
mononematous conidiophores usually unbranched or with a few branches
originating after conidial secession or near percurrent proliferations (Castaneda
etal. 1999, Taylor etal. 2001). Heteroconium avilae slightly resembles H. asiaticum
R. Chaudhary et al., but that species has pale brown, 2- to 3-septate, 40-80 x
3.5-5.5 um conidiophores and conidia that are cylindrical, tapered at the ends,
pale brown, 0- to 4-septate, and 13-24 x 5-6.5 um. Hughes (2007) comments
that H. asiaticum has probable affinities with the sooty moulds because it was
described as having superficial mycelium on living leaves. After Morgan-
Jones (1976) and Castafieda et al. (1999) outlined the characters and details
of Heteroconium species, several other species were described: H. neriifoliae
Joanne E. Taylor et al., H. eucalypti Crous & M.J. Wingf., H. glutinosum (Cooke
& Harkn.) S. Hughes & J.L. Crane and H. kleinziense Crous & Z.A. Pretorius,
H. triticicola Kwasna & G.L. Bateman and Heteroconium anam. (Hughes of
Antennulariella concinna (L.R. Fraser) S. Hughes. Helminthosporium solaninum
Sacc. & P. Syd. (as “Helmisporium’) the basionym of Heteroconium solaninum
(Sacc. & P. Syd.) M.B. Ellis was recently designated as type species of the genus
Pirozynskiella S. Hughes based on “their obligate association with asterinaceous
fungi, by general rather than apical (acropetal) extension of conidium initials and
by the centrifugal (then interseptal) sequence of septation of conidia following
the first median septum” (Hughes 2007). Heteroconium chaetospira (Grove)
M.B. Ellis was transferred to Cladophialophora as C. chaetospira (Grove) Crous
& Arzanlou after a molecular study of the Herpotrichiellaceae and Venturiaceae.
Heteroconium tetracoilum (Corda) M.B. Ellis has distoseptate conidia and
was transferred to Lylea as L. tetracoila (Corda) Hol.- Jech. (as ‘tetracoilum’);
however Hughes (2007) indicated that Lylea is not an appropriate home for
this taxon, pointing out that Heteroconium tetracoilum and H. chaetospira are
both fungicolous species. The genus Heteroconium as currently circumscribed
Heteroconium (Venezuela) ... 179
probably remains polyphyletic; Hughes (2007) described the sequence of
conidium septation in H. citharexyli as “basifugal” (i.e. the first formed septum
is at the base of the conidium, with subsequent septa produced in sequence
from base to apex) and indicated that this consistent character should be
recognized and documented in the description of anamorphs. The genera
Lylea Morgan-Jones, Xenoheteroconium Bhat et al., Cladophialophora Borelli,
Septonema Corda, Phaeoblastophora Partr. & Morgan-Jones, and Taeniolella S.
Hughes are morphologically close to Heteroconium. However, Lylea produces
distoseptate conidia in unbranched chains on monoblastic, determinate,
integrated conidiogenous cells, while Xenoheteroconium forms euseptate
conidia in branched chains and two kinds of propagules that originate from
monoblastic, terminal conidiogenous cells, although sympodial proliferations
occur in fertile areas found within the conidial chains. Cladophialophora is
distinguished by micronematous or undifferentiated conidiophores (usually
reduced to conidiogenous cells) with phialidic synanamorphs often present;
pale pigmented, 0- to 1-septate, rarely (3-septate) conidia with inconspicuous
scars are produced in branched chains in the genus. Septonema is characterized
by monoblastic or polyblastic, integrated or discrete conidiogenous cells and
(0-)1- to 3-septate conidia tapering to truncate ends that are produced in
branched chains. Phaeoblastophora is distinguished by mono- or mostly
polyblastic, integrated, terminal or intercalary conidiogenous cells and
acropleurogenous, unicellular conidia produced in simple or branched chains.
Taeniolella produces semi-macronematous conidiophores that are usually
short, unbranched or sparingly branched near the base and conidia that are
2- or more septate, dark pigmented, cylindrical, and produced in acropetal,
simple or branched chains that often secede with difficulty. Heteroconium
nigroseptatum V.G. Rao, nom. nud., was never validly published (MycoBank
2008). Heteroconium queenslandicum Matsush. appears distant from the
generic concept of Heteroconium and more closely related to the genus
Parapleurotheciopsis P.M. Kirk. The species exhibits a holoblastic conidium
ontogeny and produces an acropetal branched chain through an apical
ramoconidium that secedes schizolytically, as also found in Parapleurotheciopsis
inaequiseptata (Matsush.) P.M. Kirk (the type species of Parapleurotheciopsis).
As Diploospora indica S.K. Nair & Bhat exhibits a similar conidial ontogeny,
both species could be transferred to the genus Parapleurotheciopsis, although
we propose no new combinations here.
180 ... Castafieda Ruiz & al.
Figs.11-14. Conidia of Heteroconium species, redrawn from the original descriptions.
Fig. 11. Heteroconium tropicale. Fig. 12. H. kleinziense. Fig. 13. H. ponapense. Fig. 14. H. eucalypti.
Scale is indicated by bars: Figs. 11-13= 20 um; Fig. 14 = 10 um.
reeanncnOennenet ae
ontiapiseneriaatetng
sitio rnettatae
es
es
18
7
%
Sic
Figs.15. Conidia of Heteroconium avilae, from holotype (INIFAT C06/47-1).
Figs.16-18. Conidia of Heteroconium species, redrawn from the original descriptions.
Fig. 16. H. glutinosum. Fig. 17. H. arundicum. Fig. 18. H. citharexyli.
Scale is indicated by bars: Figs. 15-16, 18 = 10 um; Fig. 17 = 40 um.
Heteroconium (Venezuela) ... 181
Figs. 19-22. Conidia of Heteroconium species, redrawn from the original descriptions.
Fig. 19. Heteroconium indicum. Fig. 20. H. lignicola. Fig. 21. H. decorosum. Fig. 22. H. asiaticum.
Scale is indicated by bars: Figs. 19-20, 22= 20 um; Fig. 21 = 10 um.
24
cian tnarmnenpeR no ries
_enqpsnestbiginnnnniainnamnnaeetecwes
4 BHR ic
agneees
oe *isocean
EOC ORINE
Figs. 23-24. Conidia of Heteroconium species redrawn from the original descriptions.
Fig. 23. H. solaninum. Fig. 24. H. neriifoliae.
Scale is indicated by bars: Fig. 23 = 10 um; Fig. 24 = 5 um.
182.
.. Castafieda Ruiz & al.
Figs. 25-26. Conidia of Heteroconium species, redrawn from the original descriptions.
Fig. 25. H. triticicola. Fig. 26. H. anam. Antenulariella concinna.
Scale is indicated by bars = 10 um.
Key to accepted Heteroconium species
1
2 (1)
3 (1)
4 (3)
Conidiaswitht one.septuimn’ ieee acts fee) Sey a eae ce 2
Conidiasrequently wilt morestaan One Seépruin:. . e/a is 3
Conidia fusiform to cylindrical, pale brown to reddish brown, smooth,
23~42.3043—10.S Ie .:..s fee ee eek ee, es ke lt H. indicum
Conidia obclavate to ellipsoid, brown, smooth, 12-24 x 4-6 um_—_-H.. ponapense*
Euseptate and distoseptate conidia both present, conidia aseptate to
multi-septate, subcylindrical to ellipsoid, verruculose, brown,
10-60; x54-8 Lung: g tee ier: ce cee eee ets oe a ett H. kleinziense
Onily euseptate conidialpresenty same. oyster eee ak ge ee tee 4
COnidia, with) HOUMIOLe (ian a CUSeD (Ale SeDta ae ert et cueing ett. eee. 5
@onidia with:more than4 cuseplate Sep ian ante eee emit es ie eae 8
* Diagnosed from pure cultures.
- : ate et ai essa
——— a
Heteroconium (Venezuela) ... 183
5 (4) Conidia 2- to 3-septate, cylindrical to fusiform, slightly tapered at the
ends, pale olivaceous brown to pale brown, smooth, 16-24 x 4—4.5 um
eT ee is BE een ee een tae,» H. solaninum
Conidia more than 4:5, tir Wide sien i ete A AAR eo ES ee 6
6 (5) Conidia 1- to 4-septate, cylindrical, uniformly brown, smooth, 13.5-32.5 x
8. 45805 Minoel J saan cA gece Sh Pd Wea rar! tetra chs ost leet Rt H. lignicola
Conidia or at least their chains not uniformly pigmented .................... vi
7 (6) Conidia with middle cells darker than end cells, 0- to 4-septate, cylindrical,
tapered at the ends, pale brown, smooth, 13-24 x 5.0-6.5 uum ..... H. asiaticum
Conidia 23-33 x 8-12 um, 1- to 3- septate, rarely 4-septate, cylindrical to
fusiform, slightly navicular, brown to dark brown, end cells very pale brown,
guttulate, smooth, with band-shaped pigmented chains .............. H. avilae
S44)eNo conidia withimore than septa tw: oe eee ee ee ne eee gree nnn 9
SOMIe COM CIA WIL IOLG that Sea ar tweet ete rtnce eta ee tern ten ein nate My
9 (8) Conidia verruculose, 10-35 x 5-7 um, 0- to 6-septate, subcylindrical to
ellipsoid, brown at the base, pale brown toward the apex .......... H. eucalypti
Conidia smooth, 10-50 x 1.5-2.5(-3) um, 1- to 6-septate, narrowly ellipsoid to
cylindrical,with oval, thick-walled scar at both ends, subhyaline to pale brown
D. ATO, SE binvresseeid ON Hide. OLA eitsed! ikea’ a. rede H. triticicola*
Conidia smooth or, if not smooth, then not more than 5 um wide .......... 10
10 (9)Conidia 20-30 x 3-5 um, rarely verrucose, 3- to 6-septate, navicular, broad
fusiform to obclavate, sometimes sigmoid or curved, smooth .... H. decorosum
Conidia mostly longer than 30 um and / or wider than5 um............... 11
11(10) Conidia 3- to 6-septate, slightly constricted at the septa, cylindrical to
fusiform, tapered at the ends, pale brown, smooth, 26-52 x 4-6 um
, PETE, (ive ue. A. SMeearsows . Dis. Magoo, TOO. H. tropicale
Conidia ellipsoid and constricted at the septa, smooth, brown to dark brown,
9-75 x 5-9 um, synanamorph sometimes present, globose, aseptate,
at first smooth and hyaline, later verrucose and pale brown to dark brown,
11 um diam., produced in sparingly branched chains ........... H. glutinosum
Conidia dimorphic, slightly constricted at the septa, shorter conidia
subglobose to ellipsoid with few or no septa, longer conidia cylindrical,
multiseptate, brown to dark brown, 7.2—60 x 5.4-8.6 um
iter. rates i) OL be eters ee: H. anam. of Antennulariella concinna
12 (8)Conidia verrucose with longitudinal striations or with creases,
1- to 14-septate, subcylindrical tapered at the ends, brown, (6-)18-25(-45) x
(SO AP4 B24 S C6) ange Ee 2d ee ON EES H. neriifoliae
GontdiassnioGOei ervey eae lL PMN! co clas «ce ee ere 13
13(12) Conidia 1- to 10-septate, cylindrical to fusiform, mid to dark brown,
Boa OX Sal). Ua cases mira tteguw cereus “beam: Fibk Tima eas H. arundicum
Conidia 1- to 10-septate, slightly constricted at the septa, cylindrical to
oblong, pale brown to brown, 10-40 x 3-7 um.................. H. citharexyli
184 ... Castafieda Ruiz & al.
Acknowledgements
We are deeply indebted to Prof. Lori M. Carris (Washington State University) and Dr
Jerry A. Cooper (Landcare Research, New Zealand) and Prof. Roland Kirschner (J.W.
Goethe University, Germany) for kindly reviewing the manuscript. We thank Ciencia
y Tecnologia para el Desarrollo (CYTED RED-XII.J) and the Cuban Ministry of
Agriculture for facilities. The author RFCR thanks Pedro Crous, Uwe Braun, Ludmila
Marvanova, Cony Decock, Jerry A. Cooper, Roland Kirschner, Walter Gams and
Antonio Hernandez-Gutiérrez, for their generous and valuable assistance with literature
not available in his library.
Literature cited
Castafieda Ruiz RE, Saikawa M, Guarro J. 1999. A new species of Heteroconium from a tropical
rainforest. Mycotaxon 71: 295-300.
Chaudhary R, Gupta C, Kamal. 1991. New species of Heteroconium, Pseudocercospora and Stenella
from India. Mycol. Res.95: 1070-1073.
Crous PW, Groenewald JZ, Wingfield MJ. 2006. Heteroconium eucalypti. Fungal Planet, no. 10.
Crous PW, Mohammed C, Glen M, Verkley GJM, Groenewald JZ. 2007a. Eucalyptus microfungi
known from culture. 3. Eucasphaeria and Sympoventuria genera nova, and new species of
Furcaspora, Harknessia, Heteroconium and Phacidiella. Fungal Diversity 25: 19-36.
Crous PW, Schubert K, Braun U, Hocking AD, Shin HD, Groenewald JZ. 2007b. Opportunistic,
human-pathogenic species in the Herpotrichiellaceae are phenotypically similar to saprobic or
phytopathogenic species in the Venturiaceae Studies in Mycology 58: 185-217.
Ellis MB. 1976. More Dematiaceous Hyphomycetes. Commonwealth Mycological Institute, Kew,
507 pp.
Holubova-Jechova, V. 1978. Lignicolous hyphomycetes from Czechoslovakia 5. Septonema,
Hormiactella, and Lylea. Folia Geobotanica et Phytotaxonomica 13: 421- 442.
Hughes SJ. 1976. Sooty moulds. Mycologia 68: 628-820.
Hughes SJ. 2007. Heteroconium and Pirozynskiella n. gen., with comments on conidium
transseptation. Mycologia 99: 693-638.
Hughes SJ, Crane JL. 2006. A new name for Torula glutinosa in Heteroconium. Mycologia 98: 141-
143.
Kirk PM. (1982). New or interesting microfungi. IV. Dematiaceous hyphomycetes from Devon.
Trans.Br. Mycol. Soc. 78(1): 55-74.
Kwasna H, Bateman GL. 2007. Heteroconium sp.nov. from roots of Triticum aestivum in the United
Kingdom. Mycologia 99: 777-785.
Matsushima T. 1989. Matsushima Mycological Memoirs 6:1-100, Kobe, published by the author.
Morgan-Jones, G. 1976. Notes on hyphomycetes. XIV. The genus Heteroconium. Mycotaxon 4:
498-503.
Mycobank. 2008. http://www.MycoBank.org.
Nair SK, Bhat DJ. 2001. Diploospora indica, a new species of hyphomycetes. Mycotaxon 80: 101-_
104.
Petrak F. 1949. Neue Hyphomyzeten-Gattungen aus Ekuador. Sydowia 3: 259-266.
Taylor JE, Crous PW, Palm ME. 2001. Foliar and stem fungal pathogens of Proteaceae in Hawaii.
Mycotaxon 78: 449-490.
MYCOTAXON
Volume 105, pp. 185-190 July-September 2008
The Entolomataceae of the Pakaraima Mountains of Guyana 2:
Inocephalus dragonosporus comb. nov.
Davip L. LARGENT?, M. CATHERINE AIME”*?,
TERRY W. HENKEL’ & TIMOTHY J. BARON?
*maime@agcenter.lsu.edu
‘Department of Biological Sciences, Humboldt State University
Arcata, CA 95521, USA
"Department of Plant Pathology and Crop Physiology, Louisiana State University
Agricultural Center, Baton Rouge, LA 70803, USA
*Department of Biological Sciences, State University of New York-
College at Cortland, New York 13045, USA
Abstract — Entoloma dragonosporum, a saprotrophic agaric, is reported for the first
time from the Pakaraima Mountains of Guyana, occurring on soil in tropical rainforests.
Based on its stellate spores, clamp connections, mycenoid habit, appressed fibrillose
pileus, and pileipellis morphology, this Singerian species is transferred from Entoloma
to Inocephalus. The species is fully described and illustrated and the new combination
Inocephalus dragonosporus is proposed.
Keywords—Agaricomycotina, Basidiomycota, Dicymbe, Guayana Highlands,
Neotropics
Introduction
Entoloma dragonosporum was first described (as Rhodophyllus dragonosporus)
on soil from the Bolivian Amazon (Singer 1965). Subsequently, it has been
reported from tropical forests in central Brazil (De Meijer 2001), on rotten
wood and on soil in northern Brazil (Horak 1982, Singer & Aguiar 1986),
and on rotten wood in northeastern Brazil (Wartchow 2006). Additional
collections have been made in Trinidad (Baroni, unpublished), Belize (Baroni,
unpublished), and in Puerto Rico (Baroni 2007). Herein, this species is reported
for the first time from the Pakaraima Mountains of Guyana. It has yet to be
collected or reported from North America.
Species of Inocephalus are recognized by their mycenoid to collybioid
basidiomata with appressed fibrillose or at least not truly glabrous pilei that
are often acutely umbonate. Microscopically Inocephalus can be differentiated
from other Entolomataceae by its basidiospores that are often, but not always,
186 ... Largent & al.
4-sided or stellate, hyphae of the subpellis that are indistinct from the hyphae of
the pileal trama, and abundant formation of clamp connections at least on the
hyphae of the pileipellis. In addition the hyphal cells of the pileal and lamellar
trama are extremely long. The genus Nolanea can easily be confused with
Inocephalus because of its mycenoid to collybioid stature and acutely umbonate
pileus. However, Nolanea is differentiated by its glabrous pileus, inflated hyphae
of the subpellis that are distinct from the hyphae of the pileal trama, lack of
clamp connections, lack of 4-sided spores, and in most species the lack of stellate
spores. Basidiomata of taxa classified into Entoloma s. str. are fleshier, typically
tricholomatoid in stature, possess abundant clamp connections, lack 4-sided
or stellate basidiospores, and possess relatively short and broad hyphae in the
pileal and lamellar trama. Largent (1994) provided a more detailed discussion
of the genus Inocephalus.
Because of its mycenoid stature, papillate umbo, large stellate spores, abundant
clamp connections, long hyphal cells in the pileal and lamellar trama, and the
hyphae of the subpellis that are not differentiated from the hyphae of the pileus
trama, Entoloma dragonosporum is a good example of an Inocephalus species,
and the new combination Inocephalus dragonosporus is here proposed.
Materials and methods
Collections were made during the mid-May to mid-June rainy seasons of 2000-04 from
the Upper Potaro River Basin, within a 5 km radius of a permanent base camp at 5°
18’ 04.8’’ N; 59° 54’ 40.4’’ W; elevation 710m. Collections were made in both mixed
tropical forest, and in forests dominated by the leguminous, ectomycorrhizal (EM)
tree Dicymbe corymbosa Spruce ex Benth. Macroscopic features were described fresh
in the field. Colors were described subjectively and coded according to Kornerup &
Wanscher (1978), with color plates noted in parentheses. Abbreviations used in specific
descriptions indicate page number, the column, and the row; for example 8D-F5-6
indicates page 8, columns D to EF, and from rows 5 to 6. Herbarium designations are
according to Holmgren et al. (1990).
Micromorphological features of dried specimens were examined with a trinocular
research-grade Nikon Labophot compound microscope fitted with light contrast optics.
General techniques used follow Largent (1994); techniques used for measuring spores of
Entolomataceae followed Baroni (1998). Digitized photomicrographs were made using a
Nikon Coolpix 990 focused through the trinocular head of the compound microscope.
All microscopic measurements were obtained using a GTCO Corporation Graphic
Digitizer, Model DP5A-111A connected to an IBM compatible Chem Book Laptop
computer. The software utilized was Measure Me 101 v 1.0 which is modified for use °
on a contemporary laptop computer from a BASIC program called Metrics5 developed
and provided by David Malloch, University of Toronto. The mathematical results from
this program are summarized in Largent (1994). Factors determined include: X, the
arithmetic means of spore length and spores width + standard deviation; E, the quotient
of spore length by spore width indicated as a range variation in n objects measured; Q,
Inocephalus dragonosporus comb. noy. (Guyana) ... 187
Figure 1: Inocephalus dragonosporus basidiomata.
a. MCA 1563; b. field aspect (MCA 968); c. papilla (MCA 1563).
the mean of E-values + standard deviations, and for spores only, L-D the length-diameter
of spores, and AL-D the average L-D + standard deviation for n spores measured; n/4
indicates the number of objects measured/in the number of collections studied.
Taxonomy
Inocephalus dragonosporus (Singer) T.J. Baroni & Largent comb. nov. FIGS 1-2
MycoBANK MB511283
= Rhodophyllus dragonosporus Singer, Atas Inst. Micol. 2: 46. 1965.
= Entoloma dragonosporum (Singer) E. Horak, Sydowia 29: 292. 1977.
PILEUS 6-27 mm tall, 7-25 mm broad at base, conic, rounded at apex and
terminated by an acute nipple-like papillus which is ephemeral with age, off-
white to hyaline at first, glabrous, 1.5-2 mm x 1 mm; surface moist, brown (6E4-
188 ... Largent & al.
4
Figure 2: Inocephalus dragonosporus micromorphological characters. a. basidiospores (MCA
1563); b. basidia (MCA 1230); c. cyanophilic basidia (TH 8663); d. cheilocystida (TH 8663).
5) to dark brown (5E6) throughout with darker brown translucent striations,
densely radially appressed-fibrillose, fibrils longitudinally striate over entire
length; pileus margin crenate to shallowly scalloped. LAMELLAE adnate to sub-
adnexed, oriented long and almost parallel, narrow to moderately broad, 0.5-4.0
mm broad at pileus margin, 1 mm centrally, 0.5-2.0 mm at stipe apex, off-white
and pinkish with age (near 5B2), sub-thick, distant; edges roughened, at times
with some intervenous connections; lamellulae 0.5 mm long. Stipe 81-200 mm
x 1.5-2.0 mm (centrally and at apex), broadening gradually to 2-7 mm at base,
off-white, slightly graying upon pressure, fragile, smooth macroscopically but
finely longitudinally fibrillose under hand lens; basal mycelium white. ODOR
none; TASTE not obtained.
SPORE PRINT dull salmon pink on stipe apex. Spores distinctly 4—-5-sided,
stellate in all views with extremely elongated corners and concave sides, large,
22.6-36.7 x 19.4-34.5 um, (X = AL 28.52 + 2.84 x 27.16 + 3.0 um, E = 0.9-1.34,
Q = 1.05 + 0.08, L-D (-)2.64-6.56 um, AL-D 1.36 + 1.92 um; n/4 =82). BAsIDIA;
broadly cylindric with parallel sides and a rounded base, 38.5-66.0 x 16.5-22.9 -
pune r=649:07 14.0 5 lO 8 Teal OS ela R34 Ge oO A Sata ()3 i] «
n/4 = 71); distinctly cyanophilic in Amman’s; 2-4 sterigmate; sterigmata small
and indistinct. LAMELLAR TRAMA HYPHAE Subparallel, some with distinct
globules as seen in Amman’s and KOH/Phloxine, broad with long hyphal cells,
Inocephalus dragonosporus comb. nov. (Guyana) ... 189
214.5-891.1 x 9.8-32.6 um. CHEILOCyYSTIDIA abundant, forming a sterile
gill edge, broadly clavate to napiform and with distinct granular contents,
colorless but staining reddish in KOH/Phloxine, 39.6-104.5 x 9.1-35.7 um.
PLEUROCYSTIDIA absent. PILEIPELLIS an entangled layer of hyphae on the
umbo area; subpellis not differentiated from the suprapellis and intergrades
imperceptibly with the pileus trama; terminal cells cylindric to cylindro-clavate,
moderately broad and long, too few to measure. PILEUS TRAMA HYPHAE with
abundant globules or granules as seen in Amman’s and in KOH/Phloxine;
broad with long hyphal cells, 192.9-427.5 x 7.9-25.9um. STIPITIPELLIS with
rare to scattered clusters of hymenial elements, otherwise a cutis; stipe trama
very broad. PIGMENTATION suggestively cytoplasmic, occasionally with minute
patches of material adhering to the inner hyphal walls of the pileipellis near the
subpellis region. CLAMP CONNECTIONS present but inconspicuous at the base
of the basidioles, basidia, and cheilocystidia, and on the pileipellis hyphae.
MATERIAL STUDIED: GUYANA. REGION 8 POTARO-SIPARUNI: | Pakaraima
Mountains, Upper Potaro River, 20 km east of Mt. Ayanganna, near confluence of Potaro
River and Alukyadongbaru Creek, Ayanganna airstrip, growing in shelter of standing
snag, 17 May 2000, Aime 968 (BRG, LSU), near Potaro base camp, 14 June 2000 Aime
1230 (BRG, LSU), Dicymbe Plot 3, on soil, 18 May 2001 Aime 1466 (BRG, LSU), Dicymbe
Plot 2, sheltered in hollow of standing snag, 22 May 2001, Aime 1563 (BRG, LSU), Mixed
Forest Plot 1, 15 June 2001 Henkel 8147 (BRG, HSU), Mixed Forest Plot 3, 8 June 2004,
Henkel 8663 (BRG, HSU).
Hasit, HABITAT AND RANGE: Solitary to scattered on leaf humus or soil,
at times sheltered at base of standing snags or under logs in mixed tropical
rainforest or in forest dominated by Dicymbe corymbosa. Known from central
and northeastern South America (Brazil, Guyana), Caribbean Central America
(Belize) and the Greater and Lesser Antilles (Puerto Rico, Trinidad).
REMARKS—Inocephalus dragonosporus can be recognized by its narrowly conic
pileus with a hyaline to white nipple that disappears with age (Fig. 1c), extremely
long and narrow stipe measuring 81-200 mm x 1.5-2 mm (Fig. 1a), unique,
very large, stellate spores with 4-5 distinctly protruding angles like the corners
of a pagoda (Fig. 2a), large, broadly cylindric basidia with nearly parallel sides
and rounded basal ends (Figs. 2b-c), clavate colorless cheilocystidia (Fig. 2d),
and small clamp connections. Heretofore, all published illustrations of these
distinctive features have been line drawings. Cheilocystidia were not mentioned
or diagrammed by Horak (1977, 1982) or by Singer (1965).
Rhodophyllus pinnus Romagn. from Madagascar is the only other species
remotely similar to I. dragonosporus and it can be recognized by its larger pileus
(30-40 x 35-40 mm), smaller basidia (32-48 x 11.5-15 um), lack of hymenial
cystidia, and much smaller spores ((12) 14-16.5 x 10-15 um).
Of the more than 50 species of Entolomataceae thus far recorded from
the Pakaraima Mountains (Largent et al. 2007, and unpublished), Entoloma
190 ... Largent & al.
viscaurantium E. Horak & Singer is the only species whose spore shape remotely
resembles that of I. dragonosporus. However, E. viscaurantium can easily be
differentiated by its viscid, orange brown pileus, lamellae that are pale pinkish
yellow at first, then yellowish-orange (4A7-8), a much shorter stipe, and smaller
spores (7.8-13.6 x 7.2-12.8 um).
Acknowledgments
Field work in Guyana was made possible by grants from the National Geographic
Society’s Committee for Research and Exploration (TWH), the Smithsonian Institution's
Biological Diversity of the Guianas Program (TWH), the Explorer’s Club, Washington
Group Exploration and Field Research Program (MCA) and a field research gift from
W.K. Smith to MCA. We are grateful to Mimi Chin, Christopher Andrew, Francino
Edmund, and Luciano Edmund for expert field assistance in Guyana. TJB acknowledges
support from the National Science Foundation for grants DEB 0103621 and DEB
9525902, and the USDA Forest Service, Center for Forest Mycology Research, Forest
Products Laboratory which provided logistical support, for making possible field
work in Belize and the Caribbean to obtain additional comparative information on
entolomatoid species. Finally, we thank Jean Lodge and Andy Methven for their reviews
of this manuscript. This paper is No. 138 in the Smithsonian Institution's Biological
Diversity of the Guiana Shield Program publication series.
Literature cited
Baroni TJ, Lodge DJ. 1998. Alboleptonia from the Greater Antilles. Mycologia 90(4): 680-696.
Baroni TJ. 2007. Basidiomycetes of the Greater Antilles. www.cortland.edu/nsf/ga.html (uploaded
on 27 April 2007).
De Meijer AAR. 2001. Mycological work in the Brazilian state of Parana. Nova Hedwigia 72 (1-2):
105=159.
Holmgren PK, Holmgren NH, Barnett LC, eds. 1990. Index herbariorum part 1. The Herbaria of
the World. 8" ed. New York: New York Botanical Gardens. 693 pp.
Horak E. 1977. Addition to “On cuboid spored species of Entoloma”. Sydowia 29: 289-299.
Horak E. 1982. Entoloma in South America II. Sydowia 35: 75-99.
Kornerup A, Wanscher JH. 1978. Methuen handbook of colour. 3" edition. Chickester, Sussex:
Richard Clay Ltd. 252 p.
Largent DL. 1994. Entolomatoid Fungi of the Western United States and Alaska. Mad River Press
Inc., Eureka, California. 495 pp.
Largent DL, Henkel TW, Aime MC, Baroni TJ. 2007. The Entolomataceae of the Pakaraima
Mountains of Guyana I. New species of Entoloma s. str. Mycologia (in press)
Singer R. 1965. Interesting and new agarics from Brazil. Atas do Instituto de Micologia da
Universidade do Recife. 2: 15-47. ;
Singer R, Aguiar IA. 1986. Litter decomposing and ectomycorrhizal basidiomycetes in an Igap6
Forest. Plant Systematics and Evolution. 153 (1-2): 107-117.
Wartchow F. 2006. The Neotropical Entoloma dragonosporum (Agaricales, Basidiomycota): New
Record from Northeast Brazil. Biociéncias, Porto Alegre, 14 (1): 93-94.
MYCOTAXON
Volume 105, pp. 191-194 July-September 2008
Reclassification of Bremia domingensis to the genus Plasmopara
as P. domingensis comb. nov.
YOUNG-JOON CHOI & HYEON-DONG SHIN*
yjchoi@korea.ac.kr & hdshin@korea.ac.kr
Division of Environmental Science and Ecological Engineering, College of Life Sciences
and Biotechnology, Korea University, Seoul 136-701, Korea
Abstract — In this study, the isotype of Bremia domingensis on Parthenium
hysterophorus is carefully investigated, described and illustrated based on morphological
characteristics. It is concluded that the species is misplaced in Bremia but is a member
of Plasmopara. Therefore, the new combination Plasmopara domingensis is proposed
for this pathogen.
Key words — branching pattern, downy mildew, sporangia, taxonomy
Introduction
The family Asteraceae is one of the largest families of dicotyledons, within
which many genera are host plants of parasitic fungi such as Albuginaceae,
Peronosporaceae, Erysiphaceae and Uredinales. Of the numerous genera
within the Peronosporaceae, Bremia and Plasmopara are most common on
the Asteraceae. In the past, the taxonomy has been based primarily on subtle
differences in the shape or dimensions of the sporangia and upon the host plant or
assumed host range. This sometimes resulted in confusion regarding identifying
the species or even the genus of a causal agent. During a taxonomic revision of
Bremia species, we examined the type material of Bremia domingensis, a downy
mildew pathogen on Parthenium hysterophorus belonging to the Asteraceae.
The morphological characters of the fungus do not agree with Bremia. A new
combination in Plasmopara is proposed here.
Materials and methods
Herbarium specimens from the collections of U.S. National Fungus Collections,
USA (BPI) were morphologically examined. For morphological analysis,
specimens were moistened with 70% alcohol and fungi were transferred to 60%
lactic acid on a slide. The microscope preparations were warmed and examined
in brightfield- and DIC- light microscopy. Measurements were performed at
1000x for sporangia and 100-1000x for other organs.
192 ... Choi & Shin
Results
Description of Bremia domingensis —- LESIONS present on leaves, poly-angular,
2-5 mm diam., vein-limited, sometimes covering larger areas by coalescing.
Down hypophyllous, whitish, consisting of agglomerated sporangiophores, felt-
like, dense. HAusToriA intracellular, globose, unbranched, symmetrical, 4-7
um diam., surrounded by sheaths of about 1 um thickness. SPORANGIOPHORES
emerging through stomata, hyaline, straight to slightly curved, slender, 300-
420(-550) um long; trunk straight to slightly curved, 140-250(-320) um
long, 8-13 um wide, basal ends not differentiated to slightly bulbous, callose
plugs sometimes present; branches arising at right angles to the main axis,
not broadening towards the ramifications, monopodial, of 4 to 6(-7) orders,
straight to slightly curved, callose plugs often present. ULTIMATE BRANCHLETS
straight to slightly curved, (6—)8-13(-17) um long, 1.5-3 um wide at the base,
callose plugs mostly absent; tip truncate or cup-like, rarely round. SPORANGIA
hyaline, broadly ellipsoidal to subglobose, uniform in size, 15-21.3 um long
(mean=18.6 um), 13.8-17.5 um wide (mean=15.7 um), length/width ratio
(1.09—)1.15-1.26 (mean=1.19), greatest width median or sub-median, tip and
base round; pedicel absent or having a slightly eccentric scar visible at the place
of attachment; apical dehiscence apparatus 3-4 um wide, 0.8-1.3 um thick.
RESTING ORGANS not seen.
Discussion
In subfamily Asteroideae of the Asteraceae, three genera with branched
sporangiophores, Bremia, Paraperonospora and Plasmopara, are known to
cause downy mildew. Plasmopara and Paraperonospora species occur on
more hosts than do Bremia species. Sporangiophores not broadening towards
the ramifications (Fig. 1A, C) distinguished the isotype material of Bremia
domingensis from Paraperonospora. Besides, in Paraperonospora the papilla is
mostly absent or observed only as reminiscent of a dehiscence apparatus in
some species (Constantinescu 1989), but it is clearly visible on the specimen
examined (Fig. 1F-H), supporting that this fungus is not to be placed in
Paraperonospora.
The most useful morphological characteristics for distinguishing between
Plasmopara and Bremia are the more pronounced monopodial branching pattern
of the sporangiophores in Plasmopara and the inflated distal end of the
sporangiophore branches, carrying the regular aggregated ultimate branchlets, -
in Bremia (Thines 2006). In Bremia domingensis, the branches and their
subdivisions branched monopodially at right angles. The uninflated ends of the
branches and the presence of a dehiscence apparatus places B. domingensis in
Plasmopara. Two new genera, Novotelnova (Voglmayr & Constantinescu 2008)
Plasmopara domingensis comb. nov. ... 193
Fig. 1. Bremia domingensis on Parthenium hysterophorus. A: Sporangiophore; B: Haustorium; C:
Branches; D & E: Ultimate branchlets; F, G & H: Sporangia. Scale bars = 100 um for A, 10um for B,
D & E, 50 um for C, 20um for F, G & H. Source: BPI 845215 (isotype).
and Protobremia (Voglmayr et al. 2004), have recently been introduced to
accommodate the pathogens on Scorzonera and Tragopogon, respectively, in the
subfamily Cichorioideae of the family Asteraceae. The present fungus differed
from the former genus by uninflated distal ends of the sporangiophores and
from the latter by sporangiophores not broadening upward.
194 ... Choi & Shin
When Ciferri (1957) described B. domingensis on Parthenium hysterophorus,
he noted that “The morphology of our specimen agrees well with B. lactucae
Regel sensu lato”, and supposed that “this fungus is specialized on the new host
plant as a new geographical race of the genus’. Later, based on the wide species
concept of the genus, Evans (1987) proposed to treat it as a synonym under
B. lactucae. In the present study, however, we show that the morphological
characters of the type material are clearly different from B. lactucae, but agree
with that of Plasmopara.
Taxonomy
Due to the findings outlined above, it is concluded that the species described as
Bremia domingensis is misplaced in the genus Bremia, but belongs to the genus
Plasmopara instead. Therefore, a new combination in Plasmopara is proposed.
Plasmopara domingensis (Cif.) Y.J. Choi & H.D. Shin, comb. nov. FIG. 1
MycoBank # MB 511835 .
Basionym: Bremia domingensis Cif., Sydowia 10: 131, 1957 (‘1956’).
SPECIMENS EXAMINED — DOMINICAN REPUBLIC: Cibao Valley, Santiago Province,
Hato del Yaque., in living leaves of Parthenium hysterophorus L., 1930-1931, R. Ciferri
(R. Ciferri Micoflora Domingensis Exsiccata 357, holotype; BPI 845215 & BPI 786230,
isotypes).
Known distribution: only from the type collection.
Acknowledgments
The authors are grateful to James Cunnington and Marco Thines for critically reading
the manuscript and serving as pre-submission reviewers and to the curator of BPI (U.S.
National Fungus Collections, USA) for allowing access to specimens in their care. Part
of this work was supported by 2007 year’s grant from Korea University.
Literature cited
Ciferri R. 1957 (‘1956’). Micoflora domingensis exsiccata (Cent. IV, No. 301-425). Sydowia 10:
130-180.
Constantinescu O. 1989. Peronospora complex on Compositae. Sydowia 41: 79-107.
Evans HC. 1987. Fungal pathogens of some subtropical and tropical weeds and the possibilities for
biological control. Biocontrol News and Information 8: 7-30.
Thines M. 2006. Evaluation of characters available from herbarium vouchers for the phylogeny
of the downy mildew genera (Chromista, Peronosporales), with focus on scanning electron
microscopy. Mycotaxon 97: 195-218. ;
Voglmayr H, Constantinescu O. 2008. Revision and reclassification of three Plasmopara species
based on morphological and molecular phylogenetic data. Mycol. Res. 112: 487-501.
Voglmayr H, Riethmiller A, Gdker M, Weiss M, Oberwinkler FE. 2004. Phylogenetic relationships
of Plasmopara, Bremia and other genera of downy mildew pathogens with pyriform haustoria
based on Bayesian analysis of partial LSU rDNA sequence data. Mycol. Res. 108: 1011-1024.
MY COTAXON
Volume 105, pp. 195-202 July-September 2008
Wardomycopsis litoralis,
a new soil-borne hyphomycete from Spain
CAROLINA SILVERA-SIMON, JOSEPA GENE”,
JOSEP CANO & JOSEP GUARRO
carolina. silvera@estudiants.urv.cat, josepa.gene@urv.cat,
josep.cano@urv.cat, josep.guarro@urv.cat
Unitat de Microbiologia, Facultat de Medicina i Ciéncies de la Salut,
Universitat Rovira i Virgili, 43201-Reus, Tarragona, Spain
Abstract — The new hyphomycete Wardomycopsis litoralis, isolated from a soil
sample in Spain, is described and illustrated. This fungus is characterized mainly by
conidiogenous cells that show distinct terminal annellations and conidia with a median
germ slit running longitudinally. It can be differentiated from the other species of the
genus mainly by conidial morphology. Sequence analysis of the internal transcribed
spacer regions and the 5.8S ribosomal RNA gene confirmed this hypothesis and
revealed a close genetic relationship of W. litoralis with the genera Wardomyces and
Scopulariopsis.
Key words — soil fungi, conidial fungi, molecular phylogeny.
Introduction
During a continued survey of soil-borne microfungi from different
geographical areas of Spain, we recovered several interesting fungi from the
Mediterranean region. One of these was considered to belong to the anamorph
genus Wardomycopsis Udagawa & Furuya, but its morphological features did
not match those of the three other species of the genus (Udagawa & Furuya
1978, Taylor & Hyde 2003). This fungus is therefore proposed here as a new
species. To confirm our hypothesis and to infer the phylogenetic relationships
between the new fungus and other species of Wardomycopsis and other related
genera, we compared the sequence of internal transcribed spacer (ITS) regions
including the 5.8S ribosomal RNA (rRNA) gene.
Materials and methods
The samples were recovered from beach soil from Vinards, Castell6 (Spain). This
geographical area has an altitude of 35 m, an average annual temperature of 10.4—25°C
196 ... Silvera-Simén & al.
and an average annual rainfall of 12-62 mm. Soil samples free of organic matter (A,
horizon) were introduced separately into sterilized polyethylene bags, closed with rubber
bands and labelled. In the laboratory, they were stored in a refrigerator at 4—7°C until
they were processed. Soil suspensions were cultured on the selective medium Dichloran
Rose-Bengal chloramphenicol agar (Oxoid, United Kingdom) with cycloheximide (10
g/mL) and incubated at 23-25 °C. In order to study macro- and microscopic features,
the developed fungi were transferred, using a sterile dissection needle, to three different
media, i.e. potato carrot agar (PCA; 20 g potatoes, 20 g carrots, 20 g agar, 1 L sterile
distilled water, home-made), oat meal agar (OA; 30 g flakes, 20 g agar, 1 L sterile distilled
water, home-made) and potato dextrose agar (PDA; Difco Laboratories, Detroit,
Michigan), and incubated at 25°C in darkness. Growth temperature was determined
on PDA at 25, 37 and 40°C. For the observations to the optical microscope the fungal
isolates were mounted in lactic acid. Micrographs were obtained with a Leitz Dialux 20
EB microscope and a Jeol JSM-6400 scanning electron microscope.
Molecular study
The strains included in the phylogenetic study are shown in Table 1. The ITS regions
and 5.8 rRNA gene were sequenced using ITS-5 and ITS-4 primers. DNA extraction,
amplification and sequencing were performed as described in Solé et al. (2002). The
sequences were aligned using the Clustal X (version 1.81) computer programme for
multiple sequence alignment (Thompson et al. 1997). Cladistic analyses using the
neighbour-joining method (Saitou & Nei 1987) were performed with the MEGA 2.1
computer program (Kumar et al. 2001). The tree was constructed using the Kimura-2-
parameter distance model (Kimura 1980) with pair-wise deletion of gaps. The robustness
of branches was assessed by bootstrap analysis with 1000 replicates. The tree was rooted
with a sequence of Petriella setifera (Alf. Schmidt) Curzi obtained from the GenBank.
Table 1. List of species included in the molecular analysis
SPECIES STRAIN SOURCE EMBL ACCESSION NO.
Microascus trigonosporus CBS 665.71 Soil, USA AM774156*
Scopulariopsis brevicaulis a--- F tyes AJ853776
Scopulariopsis chartarum FMR 3997 Soil, Spain AM774160*
“-- --- AY625066
Wardomyces humicola CBS 369.62 (T) Soil, Canada AM774157*
Wardomyces inflatus CBS 216.61 (T) Soil, Canada AM774155*
Wardomycopsis humicola FMR 3993 Soil, Spain AM774159*
Wardomycopsis litoralis CBS 119740 (T) Soil, Spain AM774158*
Petriella setifera --- --- AF043596
CBS, Centraalbureau voor Schimmelcultures, Utrecht, The Netherlands; FMR, Facultat de Medicina de Reus,
Spain; T, ex-type culture; *, newly generated sequences for this study.
a ———
Wardomycopsis littoralis sp. nov. (Spain) ... 197
Taxonomic description
Wardomycopsis litoralis Silvera, Gené, Cano & Guarro, sp. nov. Figs 1, 2
MycoBaANK MB511237
Ad fungos conidiales, hyphomycetes pertinens. Coloniae in vitro restrictae, fuscae.
Conidiophora semi-macronemata, septata, laevia, hyalina vel subhyalina, simplicia
vel irregulariter ramosa, up to 20 um longa. Cellulae conidiogenae hyalinae vel leviter
coloratae, laeves, ampulliformes, 5.5-7.5 x 3.5-5 um. Conidia sicca, acrogena,
plerumque breve catenulata, unicellularia, obovoidea vel late ellipsoidea, 5—7 x 3—-4.5
um, primum pallida, deinde olivaceo-brunnea, crassitunicata, laeves, cum fisura mediana.
Teleomorphosis ignota.
Holotype: Spain, Catalunya, Castell6, Vinards: isolated from beach soil, Jul. 2005, J.
Gené and A.M. Stchigel (IMI 394093). Cultures ex type: CBS 119740 and FMR 8876.
EryMoLoey: from the Latin litoralis, referring to the geographical region where the
fungus was collected.
Conidial fungi, hyphomycetes. COLONIES in vitro restricted, cottony to powdery,
pale to dark grey. MYCELIUM sparse, superficial or immersed, hyaline to pale
yellowish brown, composed of mostly flexuous, branched,septate, smooth-
walled hyphae, 1.5-3 um diam, often forming funicles. CONIDIAL STRUCTURES
as annellidic conidiogenous cells borne singly along the aerial or prostrate
hyphae, or in small divergent groups on short conidiophores. CONIDIOPHORES
semi-macronematous, mononematous, septate, smooth-walled, hyaline to
subhyaline, simple or branched, up to 20 um long. CONIDIOGENOUS CELLS
Fig. 1. Wardomycopsis litoralis, CBS 119740. Conidiophores (arrows), conidiogenous cells
(arrowheads) and conidia with germ slit. Scale bar: 10 um.
198 ... Silvera-Simon & al.
Fig. 2. Wardomycopsis litoralis, CBS 119740. A, B. Conidiophores (white arrowheads), conidiogenous
cells (black arrowheads) and conidia with germ slit (arrows). C. Detail of a conidia with germ slit
(arrow). Scale bars: A-C = 10 um.
hyaline to subhyaline, smooth-walled, ampulliform, 5.5-7.5 x 3.5-5 um,
with a narrow terminal neck formed by successive annellations. Conip1A dry,
acrogenous, borne in basipetal succession as a curved short chain up to 4-5
conidia, obovoid or broadly ellipsoidal, 5-7 x 3-4.5 um, truncate at the base,
subhyaline when young, dark olivaceous when mature, thick- and smooth-
walled, with a median longitudinal germ slit. Teleomorph unknown. ;
CULTURAL CHARACTERISTICS: The optimal growth was at 25°C. At this
temperature and after 14 days of incubation, colonies on OA were 14-25 mm
diam, cottony at centre to powdery at outwards, dark grey, margin diffuse;
reverse colourless. Colonies on PCA were 17-19 mm diam, powdery, dark
Wardomycopsis littoralis sp. nov. (Spain) ... 199
grey; reverse colourless. Colonies on PDA were 13-16 mm diam, velvety,
raised, slightly fasciculate at the centre, radially folded, white at first, becoming
pale grey, margin slightly lobed; reverse colourless. At 37°C the fungus grew
more slowly than at 25°C, with colonies attaining a diameter of 6-10 mm at 14
days. The fungus was unable to grow at 40°C. The best sporulation was on OA
and PCA at 25°C.
ADDITIONAL SPECIMENS EXAMINED: Wardomycopsis humicola. Canada, Ontario,
Guelph: from soil, Aug. 1966, G.L. Barron (CBS 487.66 type strain). Spain, Catalunya,
Girona, Pals: from river sediments of the River Ter, Jun. 1991, J. Gené and K. Ulfig (FMR
3993).
Discussion
Wardomycopsis is one of the anamorph genera related to Microascaceae
(Malloch 1970, von Arx et al. 1988). This genus is characterized by dark,
globose, thick-walled conidia with germ slits that form short chains on
annellidic conidiogenous cells (Udagawa & Furuya 1978). Wardomycopsis
shows morphological similarities with Wardomyces ET. Brooks & Hansf. and
Scopulariopsis Bainier, but the members of Wardomyces produce single conidia
with germ slits from polyblastic conidiogenous cells and those of Scopulariopsis
produce long catenate conidia without germ slits (Domsch et al. 1980). Limits
between Wardomycopsis and Scopulariopsis are still unclear (Barron 1966;
Domsch et al. 1980).
The other currently accepted Wardomycopsis species are the type species —
W. inopinata Udagawa & Furuya — and W. humicola (G.L. Barron) Udagawa &
Furuya and W. trachycarpicola Joanne E. Taylor et al. (Udagawa & Furuya 1978,
Taylor & Hyde 2003). They differ from W. litoralis mainly by their conidial
morphology. W. trachycarpicola also differs from the other Wardomycopsis
species by the absence of conidial chains. In addition, we found that W. humicola,
which otherwise resembles W. litoralis morphologically, was unable to grow at
37°C. The diagnostic morphological features of the four Wardomycopsis species
are keyed out below.
Figure 3 shows the phylogenetic tree inferred from the analysis of the ITS
sequence of Wardomycopsis strains and species in other genera morphologically
close to Wardomycopsis. Unfortunately, W. inopinata and W. trachycarpicola are
not included in this phylogenetic analysis because neither strain is available in
any reference culture collections. Although the W. humicola type strain was
examined morphologically and found to be identical to the protologue and
strain FMR 3993, its sequence is considerably shorter (ca. 300 bp) than the rest
and so also could not be included in the tree. However, the type strain sequence
was informative enough to show that it was genetically very close to FMR 3993,
the isolate of W. humicola that is included in the analysis. The tree topology
200 ... Silvera-Simon & al.
supports W. litoralis as an independent species. Three main clades were clearly
distinguished in the tree. The first clade (with 87% bootstrap support) includes
the strains of Scopulariopsis chartarum (G. Sm.) EJ. Morton & G.Sm., Microascus
trigonosporus C.W. Emmons & B.O. Dodge (also presenting a Scopulariopsis
anamorph), and Scopulariopsis brevicaulis (Sacc.) Bainier. The second clade
(99% bootstrap value) groups the two Wardomyces species, i.e., Wa. humicola
Hennebert & G.L. Barron and Wa. inflatus (Marchal) Hennebert. The third
clade (with 83% bootstrap support) includes two Wardomycopsis strains — one
corresponding to W. humicola and the other to W. litoralis — although they
appear fairly distant from each other. |
Although we think Wardomycopsis litoralis is properly characterized and
clearly justified by evident morphological traits, molecular analysis only
partially supports the new taxon. This may be due to the fact that only few
strains were included in the analysis and that the ITS region is not the most
appropriate marker for inferring phylogenetic differences, as has been noted
for other genera such as Aspergillus P. Micheli ex Link (Balajee et al. 2007),
Pseudallescheria Negr. & I. Fisch. (Gilgado et al. 2005), and Phaeoacremonium
W. Gams, Crous & M.J. Wingf. (Mostert et al. 2005). The lack of available
7 Scopulariopsis charfarum AM774160°
100 Scopulariopsis charfarum AY625066
87 Microascus trigonosporus AM774156*
72 Scopulariopsis brevicaulis AJ853776
99 Wardomyces inflatus AM774155*
Wardomyces hurnicola AM774157*
94 Wardomycopsis hurnicola AM774159°*
Wardomycopsis flitoralis AM774158°
Petriella setifera AF043596
co
0.02
Fig. 3. Neighbour-joining tree based on Kimura 2-p corrected nucleotide distances among the
ribosomal internal transcribed spacer (ITS) regions and 5.8S rDNA sequences of the nine strains
listed in Table 1. Branch lengths are proportional to distance. Bootstrap replication frequencies
(1000 replications) are indicated at the internodes.* Newly generated sequences in this study.
Wardomycopsis littoralis sp. nov. (Spain) ... 201
reference strains of the species studied here constitutes an important limitation
to performing a wider study. In order to establish more clearly the boundaries
among these genera, we are currently designing primers to amplify other genes
that could be more phylogenetically informative.
Key to species of Wardomycopsis
Vere tr CODIGIarMOWMILLC HANS ep aon eat ceent ce erro. Liens Scepre ees W. trachycarpicola
i Dee OL CAST SHOT CHAINS, 1 se ae eemer nS a0, 2. SON RIA eNL neh etn eRe ee 2
2a. Conidia globose to subglobose, 4-5 um diam.................... W. inopinata
mV meme OLNICIAROUCE WISCie oy, Benet ne ere ees tick BEM aes de Pn I ee 3
3a. Conidia ellipsoidal to short cylindrical, 4-5 x 2.5-3 um. No growth at 37°C
wou Sole ga Oe NE eae PE A Ss PR see a IIR aie es ae eT W. humicola
3b. | Conidia obovoid to broadly ellipsoidal, 5-7 x 3-4.5 um. Growth at 37°C
ee NC Re ee ess See ee ne. eRe ry hme hn a tera AR W. litoralis
Acknowledgements
This study was supported by the Ministry of Science and Innovation of Spain, grant
CGL 2005-07394/BOS. We are indebted to Margarita Duefias (Real Jardin Botanico de
Madrid, Spain) and Julio Mena-Portales (Instituto de Ecologia y Sistematica, La Habana,
Cuba) for their critical comments on this manuscript.
Literature cited
Arx JA von, Figueras MJ, Guarro J. 1988. Sordariaceous ascomycetes without ascospore ejaculation.
Nova Hedwigia 94: 1-104.
Balajee SA, Houbraken J, Verweij PE, Hong S-B, Yaghuchi T, Varga J, Samson RA. 2007. Aspergillus
species identification in the clinical setting. Studies in Mycology 59: 39-46.
Barron GL. 1966. A new species of Scopularipsis from soil. Antonie van Leeuwenhoek 32: 293-
298.
Domsch KH, Gams W, Anderson TH. 1980. Compendium of soil fungi. Academic Press, London-
San Francisco.
Gilgado F, Cano J, Gené J, Guarro J. 2005. Molecular phylogeny of the Pseudallescheria boydii species
complex: proposal of two new species. Journal of Clinical Microbiology 43: 4930-4942.
Kimura M. 1980. A simple method for estimating evolutionary rates of base substitutions though
comparative studies of nucleotide sequences. Journal of Molecular Evolution 16: 111-120.
Kumar S, Tamura K, Jakobsen IB, Nei M. 2001. MEGA 2: molecular evolutionary genetics analysis
software. Bioinformatics 17: 1244-1245.
Malloch D. 1970. New concepts in the Microascaceae illustrated by two new species. Mycologia
62: 727-740.
Mostert L, Groenewald JZ, Summerbell RC, Robert V, Sutton DA, Padhye AA, Crous PW. 2005.
Species of Phaeoacremonium associated with infections in humans and environmental
reservoirs in infected woody plants. Journal of Clinical Microbiology 43:1752-1767.
Saitou N, Nei M. 1987. The neighbour-joining method: a new method for reconstructing
phylogenetic trees. Molecular Biology and Evolution 4: 406-425.
202 ... Silvera-Simon & al.
Solé M, Cano J, Pitarch LB, Stchigel AM, Guarro J. 2002. Molecular phylogeny of Gymnoascus and
related genera. Studies in Mycology 47: 141-152.
Taylor JE, Hyde KD. 2003. Microfungi of tropical and temperate palms. Fungal Diversity Press.
Hong Kong. |
Thompson JD, Gibson T], Plewniak FE Jeanmongin F Higgins DG. 1997. CLUSTAL X Windows
interface: flexible strategies for multiple sequence alignment aided by quality analysis tools.
Nucleic Acids Research 24: 4876-4882.
Udagawa S, Furuya K. 1978. A new species of Microascus and its peculiar conidial state. Mycotaxon
7291-96,
MYCOTAXON
Volume 105, pp. 203-206 July-September 2008
Llimoniella muralicola sp. nov. (Ascomycota, Helotiaceae) on
Protoparmeliopsis muralis from western Turkey
MEHMET GOKHAN HALICI
‘mghalici@erciyes.edu.tr
Erciyes Universitesi, Fen Edebiyat Fakiiltesi, Biyoloji Boliimii
38039 Kayseri, TURKEY
Abstract — Llimoniella muralicola is described as new from the areoles of
Protoparmeliopsis muralis in western Turkey. It differs from the other members of
the genus in the shorter and relatively narrow ascospores. The new species is the first
Llimoniella species reported on Protoparmeliopsis muralis.
Key words — biodiversity, lichenicolous fungi, lichens
Introduction
The genus Llimoniella was originally introduced for two closely related
species, L. adnata and L. scabridula, growing as parasites in epigeous lichen
communities over gypsaceous soils (Hafellner & Navarro-Rosinés 1993). Some
of the subsequent species, which were described under Llimoniella such as L.
fuscoatrae, L. neglecta, L. pubescens and L. stereocaulorum, were transferred
to Rhymbocarpus and L. groenlandiae was transferred to Unguiculariopsis by
Diederich & Etayo (2000).
After the synopsis of the genera Skyttea, Llimoniella and Rhymbocarpus was
published by Diederich & Etayo (2000), two more species of Llimoniella were
described: L. caloplacae by Kondratyuk et al. (2006) and L. fuscatae by Hafellner
& Obermayer (2007). Here, one more species of Llimoniella is described on
Protoparmeliopsis muralis from Turkey, raising the number of species in the
genus to eight.
Material and methods
The type material of the new species is deposited in the herbarium of Erciyes
University, Science& Art Faculty, Biology Department, Kayseri. Specimens were
examined with an Olympus BH-2 research microscope fitted with Nomarski
differential interference contrast optics and a drawing tube. Photomicrographs
204 ... Halici
were prepared on a Nikon Eclipse 80i. Sections were prepared by hand and
examined in I (Lugol's iodine, with [K/I] and without [I] pre-treatment with
10% KOH), 10% KOH alone, and water. The exciple structure was examined in
C (commercial bleach). Ascospore measurements were made in water and 10%
KOH; the extreme values outside the main range are given in parentheses. The
length/breadth (1/b) ratio of the ascospores is given in the same way.
The species
Llimoniella muralicola Halici, sp. nov. - FIGURE 1
MycoBank MB 511866
Llimoniella species ascomatibus superficialibus, 0.3-0.6 mm diam., nigra, margine
prominenti. Asci unitunicati, octospori, 28-32 x 6-7 ym, parietibus apicaliter non
incrassates, non amyloideis. Ascosporae hyalinae, unicellulares, ellipsoideae, 7-8(-9)
x (2.5-)3-4(-5) um magnae, non halonatae. Habitat supra thalium Protoparmeliae
muralae.
Typus: Turkey, Denizli, Dikmen Village, Babadag, 38°43’N, 28°52’E, alt. 2065 m, on
thallus of Protoparmeliopsis muralis on siliceous rocks, 14 June 2007, leg. M. Kocakaya
(MGH 0.5268 — holotypus).
EtTyMo_ocy: The epithet “muralicola” refers to the host Protoparmeliopsis muralis.
DESCRIPTION: Lichenicolous, on the thallus of Protoparmeliopsis muralis,
causing slight discolorations in the infected areoles, mildly pathogenic.
Ascomarta at first perithecioid, gradually opening at the apex and becoming
apothecioid, dispersed on the host thallus, sometimes loosely aggregated in
groups of 2-4, superficial, matt black, initially + closed, soon applanate, 0.3-0.6
mm in diam., concave to slightly convex, margin prominent, scabrous. Exciple
persistent, with a dark reddish brown pigment, K + violet-red, 30-50 um thick;
composed of + isodiametric cells of 4-6 um diam., except the outer layer which
consists of radiating hyaline cells reaching to 20 um in length. Subhymenium
hyaline. Hymenium colourless, without a greenish tinge, K -, 40-50 um tall,
KI -, no distinct epithecium and hypothecium developed. Paraphyses hyaline,
c. 1.5-2.5 mm thick, occasionally branched, easily separating, apically slightly
thicker, 2.5-3.5 um, septate. Asci elongate-clavate, very shortly stalked, apically
not thickened, rarely hardly thickened, 8-spored, KI, I -, 28-59 x 6-9 um.
Ascosports irregularly biseriate or uniseriately arranged in the asci, ellipsoid,
hyaline, straight or curved, non-septate, generally 2-guttulate, sometimes
Pralnouttaete smooth-walled, 7-8(-9) x (2.5-)3-4(-5) um (n = 32), I/b =
(2—)2.35-2.7(-3.2). CONIDIOMATA not observed.
ECOLOGY AND DISTRIBUTION: The species appears to be mildly pathogenic as
noted from discolorations in the infected areoles of the host. The new species is
known from one locality from the western part of Turkey. As the host species has
a wide distribution in the world, the species should be searched for elsewhere.
Llimoniella muralicola sp. noy. (Turkey) ... 205
i \ A (00 0 6€
i 660008 |
en
2
Fig. 1. Llimoniella muralicola (holotype).
A, Exciple; B, Asci and paraphyses; C, Ascospores
Scale = 10 um
OBSERVATIONS: Llimoniella muralicola is the first species of Llimoniella reported
on Protoparmeliopsis muralis. The new species seems to be most closely related
to L. scabridula (Miull.Arg.) Nav.-Ros. & Hafellner, which occurs on terricolous
Acarospora species (Hafellner & Navarro-Rosinés 1993). However, L. muralicola
differs from this species in having shorter and narrower ascospores [7-8(-9) x
(2.5-)3-4(-5) um vs. (8-)9-11 x 4-5(-6) um]. L. adnata Hafellner & Nav.-
Ros., known from terricolous Placidium squamulosum (Hafellner & Navarro-
Rosinés 1993), also has longer and wider ascospores [9-13 x (5-)6-7 um vs.
7-8(-9) x (2.5-)3-4(-5) um] and paraphyses that are pale blue pigmented
in the end cells. L. fuscatae Hafellner & Obermayer, a recently described
species on Acarospora fuscata from Austria (Hafellner & Obermayer 2007) has
obviously larger and wider ascospores [9-12 x 5.5-7.5 um vs. 7-8(-9) x (2.5-
)3-4(-5) um]. Another recently described species, L. caloplacae S.Y.Kondr. &
Khodos. on Caloplaca borysthenica from Ukraine (Kondratyuk et al. 2006) has
0-1-septate ascospores which are obviously longer [9.5-13(-16) x 2.85-3.5(-4)
tum vs. 7-8(-9) x (2.5-)3-4(-5) um].
Acknowledgements
Javier Etayo and Pere Navarro-Rosinés are thanked for reviewing this paper. Alper T.
Halici and Gékhan Hinisli are thanked for their help in preparing the figures. Mustafa
Kocakaya is thanked for providing the specimens.
Literature cited
Diederich P, Etayo J. 2000. A synopsis of the genera Skyttea, Llimoniella and Rhymbocarpus
(Lichenicolous Ascomycota, Leotiales). Lichenologist 32: 423-485.
206 ... Halici
Hafellner J, Navarro-Rosinés P. 1993. Llimoniella gen.nov.- eine weitere Gattung lichenicoler
Discomyceten (Ascomycotina, Leotiales). Herzogia 9: 769-778.
Hafellner J, Oberniayer W. 2007. Flechten und lichenicole Pilze im Gebiet der Stubalpe (Osterreich:
Steiermark und Karnten). Mitt. naturwiss. Ver. Steiermark 136: 5-59.
Kondratyuk S$, Khodosovtsev A, Karnefelt I. 2006. Llimoniella caloplacae sp. nova (Leothiales),
a new lichenicolous fungus on Caloplaca borysthenica sp. nova (Lecanorales, Ascomycotina).
Mycologica Balcanica 3: 95-98.
MYCOTAXON
Volume 105, pp. 207-224 July-September 2008
Cercosporoid hyphomycetes on hosts of the Annonaceae:
Cercospora annonaceae and Isariopsis annonarum revisited
UwE BRAUN! & PEDRO W. Crous?
uwe. braun@botanik.uni-halle.de
'Martin-Luther-University, Institute of Biology, Geobotany and Botanical Garden
Herbarium, Neuwerk 21, D-06099 Halle (Saale), Germany
p.crous@cbs.knaw.nl
°CBS, Fungal Biodiversity Centre
PO. Box 85167, 3508 AD Utrecht, The Netherlands
Abstract — Type material of Cercospora annonaceae and several collections assigned to
this species by C. Chupp have been re-examined. ‘The status of this species as a member
of the genus Stenella is confirmed, but most collections in Chupp‘s herbarium referred
to as C. annonaceae proved to be misidentified, and pertain to Isariopsis annonarum,
herein reallocated to Pseudocercospora, and the new species Phacellium annonae-
cherimoliae. A key to 23 cercosporoid hyphomycete species on hosts belonging to the
Annonaceae is provided, and the taxonomy of the species concerned is briefly discussed.
The following new combinations are proposed: Pseudocercospora annonarum,
P. asiminae and P. oblecta. Furthermore, a revised, detailed description of
Pseudocercospora annonae-squamosae is provided.
Key words — anamorphs, cercospora-like, Annona
Introduction
Cercospora annonaceae (Hennings 1909) was introduced for a cercosporoid
hyphomycete on an unknown member of the Annonaceae collected in Brazil.
The original data are rather scanty, simply describing hypophyllous caespituli,
fasciculate ‘hyphae’ (i.e. conidiophores), up to 150 x 3.5 um, septate, and
cylindrical-fusiform conidia, 15-35 x 2.5-3.5 um, 3-7-septate, brown. In
his monograph of Cercospora Fresen., Chupp (1954) considered this species
a coremioid hyphomycete, which has to be excluded from the latter genus.
His observations were based on a revision of type material, deposited at the
herbarium of the Botanical Garden and Museum in Berlin, Germany (B),
and several secondary samples in his own herbarium, now maintained at the
Plant Pathology Herbarium of the Cornell University, Ithaca, New York, USA
(CUP). Unfortunately the type material at B has been lost. Braun (in Braun
208 ... Braun & Crous
& Melnik 1996) detected and examined syntype material of this species
deposited at the Russian herbarium LEP. Hennings (1908) cited the following
details for the type collection: ‘Agua Branca in foliis Anonaceae. Majo 1903,
no. 738’ [‘Agua Branca, Sao Paulo, Anonaceae, A. Puttemans, No. 738, May
10- (162), 1903’ is the altered version given in Chupp (1954), who examined
this material]. The label of the syntype material at LEP deviates and reads
as follows: ‘Para, Annonaceae, V 1903, Puttemans. The whole context of the
protologue and the labels concerned suggest the locality Agua Branca (or Agua
Branca do Paulinho?) in the state of Para, in which also a city with the name
Sao Paulo exists. Braun (Braun & Melnik 1996) reallocated C. annonaceae to
Stenella Syd., since the only discernable cercosporoid hyphomycete found in
the syntype material at LEP was a member of the latter genus. However, in the
latter material only solitary conidiophores arising from superficial hyphae were
found, i.e. fasciculate conidiophores were lacking, which was in conflict with
the original description (Hennings 1908) as well as Chupp’s (1954) comments
on this species, and raised the question whether a single polymorphous or two
different cercosporoid taxa had been involved. Therefore, herbarium specimens _
referred to as C. annonaceae by C. Chupp have been re-examined in order to
clarify his taxonomic decision and concept.
Materials and methods
The collections examined were described, mounted in distilled water, using
oil immersion (bright field and phase contrast), but without any staining, by
means of standard light microscopy (Olympus BX 50, Hamburg, Germany).
30 measurements (x 1000 magnification) of conidia and other structures were
made, with the extremes given in parentheses. The collections examined are
deposited at the herbaria CUP and W (abbreviations according to Holmgren
et al. 1990).
Taxonomy
The examination of four specimens from CUP referred to as C. annonaceae by
C. Chupp revealed them to represent several distinct taxa. A single collection
from Brazil on Annona reticulata proved to be a member of Stenella conspecific
with C. annonaceae, a specimen from Venezuela on A. squamosa belongs to
Isariopsis annonarum [= Passalora annonarum], and another one from Mexico
on A. cherimolia has to be described as new species of Phacellium Bonord.
Stenella annonaceae (Henn.) U. Braun, in Braun & Mel'nik,
Mikol. Fitopatol. 30(4): 3, 1996. Fic. 1
= Cercospora annonaceae Henn., Hedwigia 48: 18, 1908 [*1909’].
Cercosporoids on Annonaceae ... 209
MATERIAL EXAMINED: BRAZIL. Para, on living leaves of a host belonging to the
Annonaceae (probably Annona sp.), May 1903, Puttemans (LEP), lectotype of C.
annonaceae, designated here. BRAZIL. Fungos de Minas Geraes, Vicosa-Escola, on
living leaves of Annona reticulata, 23 Mar. 1930, A.S. Muller (CUP-MG 157).
LEAF SPOTS amphigenous, subcircular to angular-irregular, 1-25 mm diam.,
sometimes confluent, pale to dull dark brown, later grayish to dingy grayish
white, margin indefinite or with a narrow darker border or marginal line.
COLONIES amphigenous, mainly hypophyllous, inconspicuous, effuse to
punctiform, brown to grayish brown. MycELium internal and external;
superficial hyphae emerging through stomata, 1.5-3 um wide, branched, septate,
subhyaline, pale to medium olivaceous, olivaceous-brown or yellowish brown,
thin-walled, verruculose. Srromata lacking or small, not very conspicuous,
10-40 um diam., immersed, brown. CONIDIOPHORES solitary, arising from
superficial hyphae or in small to moderately large fascicles, loose to usually
rather dense, occasionally even coremioid, arising from immersed stromatic
hyphal aggregations, erect, unbranched, subcylindrical, straight to somewhat
flexuous-sinuous, upper fertile part usually geniculate-sinuous, sometimes
strongly so, 20-150 x (2.5-)3-4(-4.5) um, pluriseptate, pale to dark brown,
yellowish brown or olivaceous-brown, smooth, wall thin to slightly thickened
(< 1 um). CONIDIOGENOUS CELLS integrated, terminal and intercalary, 5-20 um
long, occasionally longer, with several conspicuous conidiogenous loci, slightly
thickened and darkened, 1-1.5 um diam. Conip1a solitary, subcylindrical-
filiform, acicular to narrowly obclavate, 20-100 x 2.5-4.5 um, 2-8-septate, pale
olivaceous, yellowish brown to brown, verruculose, wall thin (ca. 0.5 um wide),
apex obtuse to subacute, base truncate to usually obconically truncate, 1-1.5
(—2) um wide, hila slightly thickened and darkened.
ComMMENTSs: The collection on Annona reticulata agrees well with the
original description of C. annonaceae as well as the lectotype of this species
in that solitary as well as fasciculate conidiophores are formed. Fasciculate
conidiophores were described by Hennings (1908) and observed by Chupp
(1954), whereas solitary conidiophores arising from superficial hyphae were
found by Braun & Mel'nik (1996) in the lectotype sample.
Phacellium annonae-cherimoliae U. Braun, sp. nov. Bigs
MycoBankK MB 511858.
Phacellio sessili simile, sed conidiis longioribus et angustioribus, 8-25 x 2-4(-5) um,
0-1(-2)-septatis, levibus.
MATERIAL EXAMINED: MEXICO. Jalapa, Vera Cruz, on living leaves of Annona cherimolia
(Annonaceae), 19 Jul. 1932, O.A. Plunkett 143 (CUP 60731), holotype. Isotype: CUP
39057.
LEAF SspoTs amphigenous, subcircular to angular-irregular, 1-5 mm diam.,
occasionally confluent, pale to dark brown, later grayish brown to dingy gray,
210 ... Braun & Crous
Fig. 1. Stenella annonaceae. A. Conidiophore fascicle. B. Superficial hyphae with or without solitary
conidiophores. C. Tips of conidiophores with conidiogenous cells. D. Conidia. Scale bar = 10 um
(fig. A based on CUP-MG 157, figs B—G based on type material from LEP). U. Braun del.
Cercosporoids on Annonaceae ... 211
on the lower leaf surface usually much paler, margin conspicuous, narrow, dark
brown to blackish. Myce.ium internal, forming immersed stromatic hyphal
aggregations at the base of synnemata, 20-50 um diam., brown. CONIDIOMATA
synnematous, erect, straight, occasionally slightly curved to sinuous,
subcylindrical or somewhat attenuated towards the apex, 150-400 x 10-40 um,
medium brown, composed of a firm stipe of densely appressed parallel threads
(conidiophores) and a rather compact, often poorly developed capitulum,
barely or only slightly splaying out. INDIVIDUAL CONIDIOPHORES 1-4 um wide,
pale to mid olivaceous, olivaceous-brown or brown, pluriseptate throughout,
smooth, thin-walled. CONIDIOGENOUS CELLS integrated, terminal, intercalary
and pleurogenous, 5-20 x 2-5 um, subhyaline to very pale olivaceous or
olivaceous-brown, with a single to usually several conspicuous conidiogenous
loci, 1-1.5 um diam., slightly thickened and darkened. Conip1a catenate, in
simple or occasionally branched chains, ellipsoid-ovoid, cylindrical, 8-25 x
2-4(-5) um, 0-1(-2)-septate, hyaline or subhyaline, thin-walled, smooth or
almost so, ends obtuse to somewhat attenuated, hila 1-1.5 um diam., slightly
thickened and darkened.
CoMMENTs: Due to synnematous conidiomata with integrated, terminal as
well as pleurogenous, cicatrized, hyaline or only pale conidiogenous cells, and
hyaline or subhyaline conidia formed in chains, the new cercosporoid species
on Annona cherimolia fits the concept of the hyphomycete genus Phacellium
as circumscribed by Braun (1998). Phacellium annonae-cherimoliae, the first
species of Phacellium on a host belonging to the Annonaceae, is morphologically
close to P. sessile U. Braun, but differs in having longer and narrower, 0-1(-2)-
septate, smooth conidia. P sessile, on Sanguisorba canadensis in North America,
possesses very similar synnemata with numerous pleurogenous conidiogenous
cells, but the conidia are 10-16(-18) x 4-6(-8) um, aseptate and verruculose
(Braun 1998). The South American P paspali (Syd.) U. Braun is a further
similar species, also characterized by having slender, pigmented synnemata,
often somewhat narrowed towards the apex, with little differentiated capitula
and abundant pleurogenous conidiogenous cells, but the synnemata are pale
reddish to pink and the conidia are aseptate. The type species of Phacellium,
P. inhonestum Bonord. [= P. alborosellum (Desm.) U. Braun], is, except for
the synnematous conidiomata, close to Ramularia Unger. Other species with
distinctly pigmented synnemata and conidiophores are more cercosporoid,
resembling phaeoramularia-like Passalora Fr. species, but differ in having
pleurogenous, hyaline or subhyaline conidiogenous cells and colourless conidia.
The taxonomy of this genus within the complex of cercosporoid Mycosphaerella
anamorphs is still unclear. Cultures of the type species and molecular sequence
analyses of various Phacellium species, including those with colorless and
pigmented synnemata, are not yet available. Thus, we prefer to maintain
Phacellium tentatively.
212 ... Braun & Crous
Pseudocercospora annonarum (Petr. & Cif.) U. Braun & Crous, comb. nov. Fia. 3
MycoBankK MB 511859.
Bas.: Isariopsis annonarum [‘anonarum’] Petr. & Cif., Ann. Mycol. 30: 235, 1932.
= Phaeoisariopsis annonarum (Petr. & Cif.) U. Braun, Nova Hedwigia 50: 511, 1990.
Passalora annonarum (Petr. & Cif.) U. Braun & Crous, in Crous & Braun,
Mycosphaerella and its anamorphs: 1. Names published in Cercospora and Passalora.
CBS Biodiversity Series 1: 438, 2003.
MATERIAL EXAMINED: DOMINICAN REPUBLIC. Prov. Santiago, Santiago, Valle del
Ciboa, on living leaves of Annona squamosa (Annonaceae), 22 Dec. 1930, Ekman (W,
No. 10913), type of I. annonarum. VENEZUELA. Caracas, Los Chorros, on living
leaves of Annona squamosa, Mar. 1939, Muller & Whetzl (CUP-VZ 2892), as Cercospora
annonaceae.
LEAF spots amphigenous, subcircular to angular-irregular, 1-10 mm diam.,
occasionally confluent, pale to dark brown, later grayish brown to dingy gray,
on the lower leaf surface usually much paler, margin conspicuous, dark brown
to blackish, narrow. Conip1ioMatTA hypophyllous, synnematous, scattered,
rather inconspicuous to punctiform, brown to dark brown. Myce ium internal.
STROMATA immersed, 20-60 um diam., brown. SYNNEMATA 80-250 x 20-50 um,
brown, composed of a mostly firm subcylindrical stipe of numerous, densely
appressed parallel conidiophores and a more or less loose apical capitulum,
often somewhat wider at the very base, occasionally with conidiophores in dense
fascicles, i.e. not distinctly synnematous, individual conidiophores 40-250 x 2-5
lum, terminal cells (conidiogenous cells) up to 7 um wide, olivaceous to medium
brown, pluriseptate throughout, thin-walled, smooth. CONIDIOGENOUS CELLS
integrated, terminal, rarely intercalary or pleurogenous, 10-25(-35) um long;
conidiogenous loci inconspicuous to subconspicuous by being subdenticulate
or by having an unthickened, but slightly darkened-refractive ultimate rim
(paracercospora-like), visible in front view as minute circle, 1-2 um diam.
Conrpia solitary, obclavate-cylindrical, occasionally subclavate, short conidia
sometimes ellipsoid-ovoid, straight to curved, (15-)25-70(-80) x (3.5-)4-8
(-9) um, (0—)1-7(-10)-septate, occasionally constricted at the septa, pale to
medium olivaceous or olivaceous-brown, thin-walled (ca. 0.5 um), smooth,
apex obtuse, base obconically truncate to somewhat convex, (1.5—)2-2.5(-3)
um wide, unthickened, not darkened, at most slightly refractive.
CoMMENTSs: During the course of a revision and reassessment of the genus
Phacellium (= Isariopsis Fresen.), Braun (1990) reallocated cercosporoid
species with pigmented, scolecoid conidia, including Isariopsis annonarum,
to Phaeoisariopsis Ferraris. Later it turned out that the formation of
synnematous conidiomata is of little taxonomic relevance at generic level in
cercosporoid anamorphs. Furthermore, the genus Phaeoisariopsis proved to
be heterogeneous, encompassing passalora-like species with conspicuous,
thickened and darkened conidiogenous loci as well as pseudocercospora-
like taxa with inconspicuous loci (Deighton 1990, Crous & Braun 2003).
213
Cercosporoids on Annonaceae ...
Figs 2-3. 2. Phacellium annonae-cherimoliae. 3. Pseudocercospora annonarum. A. Synnemata.
B. Conidiophores in a dense fascicle. C. Conidiogenous cells. D. Conidia (based on type material,
except for fig. B of P annonarum which is based on CUP-VZ 2892). Scale bars
del.
= 10 um. U. Braun
Hence, most species of Phaeoisariopsis were reallocated to Passalora and
respectively. Due to subconspicuous conidiogenous loci
Pseudocercospora,
but somewhat darkened-refractive, scars visible in
(ultimate rim unthickened,
214 ... Braun & Crous
front view as minute circles), Crous & Braun (2003) assigned I. annonarum
to Passalora. Based on a molecular and morphological reassessment of
Phaeoisariopsis griseola (Sacc.) Ferraris, the type species, Crous et al. (2006)
reduced the genus Phaeoisariopsis to synonymy with Pseudocercospora.
P. griseola is characterized by having conidiogenous loci ranging from being
quite inconspicuous to subconspicuous by being unthickened, but slightly
darkened-refractive, at least with regard to the ultimate rim. Since P. griseola
clustered within a subclade formed by other Pseudocercospora species, it became
clear that taxa with minutely thickened and slightly darkened conidiogenous
loci have to be placed in Pseudocercospora. The recent examination of the rich
collection of I. annonarum from Venezuela showed that this species has a
similar morphological range of the conidiogenous loci as in P. griseola. Thus,
I. annonarum is better placed in Pseudocercospora.
Key to cercosporoid hyphomycete species on hosts of the Annonaceae
Numerous cercosporoid hyphomycete species, including the taxa treated herein, have
been described on host species belonging to the Annonaceae. The following key and brief -
discussions on some of the species listed have been prepared to render the differences
between the taxa concerned palpable.
1 Gonidiophores in'distincusyanemiata’.....).--. Beet > . | BRERE wooo as oe 2
1* Conidiophores mononematous, solitary or fasciculate ..................... 3
2 Synnemata composed of a firm stipe and a + loose capitulum; conidiogenous
cells usually integrated, terminal; conidiogenous loci inconspicuous or only
ultimate rim slightly darkened-refractive (visible in front view as minute circle);
conidia solitary, pigmented, (15—)25-70(-80) x (3.5—)4-8(-9) um; on Annona
Sqanipsa ad WET) OS eee a ee. OF, TRE Pseudocercospora annonarum
2* Synnemata composed of a firm stipe and a compact, little differentiated
capitulum; conidiogenous cells frequently pleurogenous; conidiogenous loci
minute, but conspicuous, thickened and darkened; conidia catenate, hyaline or
subhyaline, 8-25 x 2—4(-5) um, 0-1(-2)-septate; on Annona cherimolia
eRe Ee) REP 2) See as gti Phacellium annonae-cherimoliae
3 Conidiogenous loci conspicuous, thickened and darkened .................. +
3* Conidiogenous loci inconspicuous, at most subconspicuous by being denticle-
like, but neither ihickenednor darkened <C ross. een eek ee oe 11
4 Superficial mycelium present; hyphae distinctly verruculose ................ 5
4* )))Soperiicial myceum' smooth ‘or lackingoeer eo Pe ed eee 6
5 Conidiophores solitary, arising from superficial hyphae, as well as fasciculate; on
Antonia spp5 Brazile ei VIET NS Mae ee ne ee Stenella annonaceae
5* Conidiophores consistently solitary, arising from superficial hyphae; on
Polyalthia suberosa, India aioe aie wee ot Jers te. oa Stenella polyalthiae
Cercosporoids on Annonaceae ... 215
6 Conidiophores fasciculate, pigmented; conidia solitary, acicular when mature,
pluriseptate, hyaline; on Annona spp. ...........0000000. Cercospora apii s. lat.
6* Conidiophores fasciculate or solitary, pigmented; conidia not acicular, solitary
and pigmented or catenate and + hyaline (Passalora) ............0..000005. 7
Z LC OTH TG Gi RSW Sap ie caeneat os orm et aa ate RR MARCIE oye timae Peer NL 2: 8
me LOTUCiaSOlt ALY grt f Tedncoe PE EEN Bettis a ce olin so s-4 eels es Slgiloabie LOE a oes 2)
7.8 Conidiophores 120-200(-250) «sm long; conidia 30-70 x 3-4 um, 2-7-septate,
subhyaline to pale brown; on Isolona cauliflora, Kenya ...... Passalora isolonae
8* Conidiophores 30-150 um long; conidia 20-40(-65) x 2-4.5 um, 0-3(-5)-
septate, hyaline or subhyaline; on Annona dioica and Xylopia grandiflora, Brazil
Ac enetesio eel ais oles lara, © rate, iat aura read cieiprron A x Passalora xylopiae
9(7) Conidiophores solitary and fasciculate, ca. 60-210 x 5.5-9 um; conidia ca.
35-130 x 7-9 um, 1-4-septate; on Miliusa sp., India
9* Conidiophores much shorter and narrower, up to 80 x 6um............... 10
10 Conidiophores solitary, arising from superficial hyphae; conidia 14.5-50 um
long, up to 4-septate; on Miliusa tomentosa, India .......... Passalora miliusae
10* Conidiophores fasciculate, emerging through stomata; conidia 25-80 um long,
Beb-sepiate; GD AMmnOnd spp, DEAL. ana ee Gee Passalora annonigena
11(3) Superficial mycelium present; conidiophores solitary as well as fasciculate,
BOlletIlcs evel sDOLOdOCal coniciOliala mates) 2 ete arene 12
11* Superfical mycelium lacking; conidiophores consistently in loose to dense,
ALIMOSUSPOLOUOCIIALASCICCSner nett oe nt, er ee eee eee ye
12. Conidia very broad, 20-80 x 4-12 um, with 1-9 transverse and occasionally
1(-2) oblique or longitudinal septa; on Asimina triloba, North America
sks ty Paced tet RE CCR ME TO RO FR Oe ee Pseudocercospora asiminae
12* Conidia much narrower, up to 7 um wide, only transversely septate......... 13
13. Conidiophores consistently solitary, arising from superficial hyphae, distinct
fascicles lacking, 12-70 x 3.5-5 um; conidia narrowly obclavate-cylindrical,
40-156 x 2.5-3.5 um, 3-13-septate; on Polyalthia suberosa, India
PRT Iee Vee. Ek ORSAY Whom BON. BE, Pseudocercospora polyalthiae
13 ee Couidiophoressolitaryasmvellas fasciculate. 2... 0a. ne ee ee 14
14 Conidia very long and wide, 50-150 x 5-7 um, densely pluriseptate (6-18), hila
2.5-3.5 um wide; conidiophores usually fasciculate, occasionally solitary, arising
from superficial hyphae; on Annona spp. and Rollinia sp., Asia, West Indies,
Gentralto SOutnpAtneriCagwuesc ts gent i ei ante, Pseudocercospora annonae
14* Conidia narrower, 1.5-6(-7) um wide, up to 10-septate, hila narrower,
De 2r Une. oss Se onoadt ah piss TLicisi der i Aeee eT Tate Eee 15
15 Conidia 25-140 x 3-6(-7) um, very pale, mostly subhyaline; on Asimina spp.,
North} America 60s. 2h. 2asaet 3B Ses Pseudocercospora asiminae-pygmaeae
15* Conidia narrower, 1.5—4 um, subhyaline to pale brown; on Annona and Rollinia
Ey 8] OS esa tri cs, ee a ee Pa RI at A POR i etl a te yaloay fic eae 16
216... Braun & Crous
16 Superficial mycelium with solitary conidiophores abundant, fascicles of
conidiophores variable, small and loose to rich and dense, occasionally in
almost sporodochial conidiomata; apex of conidia obtuse in cylindrical conidia
to subacute in obclavate conidia; on Annona spp. and Rollinia mucosa, Asia,
West Indies, Central to South America .. Pseudocercospora annonae-squamosae
16* Mycelium internal and external, but conidiophores in dense fascicles, solitary
conidiophores lacking; apex of conidia obtuse; on Annona purpurea, Brazil
Wont, Sole Wee RANE Pes ee 8 eee 1 ete Ae Pseudocercospora xenoannonicola
17(11)Conidiophores in sporodochial conidiomata; conidiogenous cells monoblastic,
percurrent, with distinct annellations; conidia rough-walled ........ TS EGS. 18
17* Conidiogenous cells monoblastic to usually polyblastic, sympodial,
annellations lacking; conidia smooth or almost so .................200000. 19
18 Conidiophores 30-50 um long, verruculose; conidia 40-120 x 4.5—-8 um,
obclavate-cylindrical; on Annona senegalensis, South Africa
GROVE Is RRA Een RS en cn ee ee Pseudocercospora oblecta
18* Conidiophores shorter, 5—20(-—30) um long, smooth; conidia narrower, 20-220 x
3.5-5 um, cylindrical-vermiform; on Annona crassifolia, Brazil
od wien Hactce henatee ts Ml cestatd ss eho ie Mc e tae ae Nc Pseudocercospora annonifolii_ .
19(17)Conidia broad, 5-8 ttm; conidiophores in small, + loose fascicles .......... 20
19* Conidia narrower, 2—5 um; conidiophores usually numerous, in + dense,
Olen Sporodochital fasciclesw. wee weet ae Rae his ce eee eee 21
20 Conidiophores fasciculate, very long, 60-220 um; conidia obclavate,
50-110 um long, base obconically truncate; on Annona senegalensis and
ANNONA SD atiCay Asia ss cn een ee eo cen ee Pseudocercospora scitula
20* Conidiophores much shorter, up to 100 um long, fasciculate, occasionally
solitary, arising from superficial hyphae; conidia broadly acicular-obclavate,
base truncate to slightly obconically truncate; on Annona spp. and Rollinia sp.,
Asia, West Indies, Central to South America........ Pseudocercospora annonae
21(19)Conidia subcylindrical-filiform to subacicular, 57.5-115 x 3.5-4.5 um,
hyalines or Wittiisd sp., Widia ste ogee ee Pseudocercospora annonacea
21* Conidia obclavate-cylindrical, narrower, 2-3 um wide, olivaceous to
OlivaceGus-DIOWD cme wah «oo. edo Cees, Stat tee a eae 22
22 Conidia (30—)50-80(—100) um long; mycelium internal and external;
on Annona montana, Brazil ................ Pseudocercospora xenoannonicola
22* Conidia shorter, 30-65 um long; mycelium consistently internal;
ORLA yODIA. GeINIOPICa: AIC. 40 san ee eee Pseudocercospora aethiopicae
Cercospora apii Fresen. s. lat. (sensu Crous & Braun 2003).
COMMENT: C. apii s. lat. (incl. C. canescens Ellis & G. Martin) has been
recorded on Annona ‘odorata’ (Crous & Braun 2003).
Cercosporoids on Annonaceae ... 217
Passalora annonacearum [‘anonacearum’ | A.N. Rai & Kamal,
Kavaka 14: 33, 1988 [‘1986’].
ILLUSTRATION: Rai & Kamal (1988: 34, Fig. 2).
On Miliusa sp., Asia, India.
Passalora annonigena U. Braun & EO. Freire,
Cryptog. Mycol. 23: 297, 2003 [‘2002’].
ILLUSTRATION: Braun & Freire (2003: 298, Fig. 2).
On Annona sp., South America, Brazil.
Passalora isolonae (Siboe, PM. Kirk & PE Cannon) U. Braun & Crous,
in Crous & Braun, Mycosphaerella and its anamorphs. 1. Names published
in Cercospora and Passalora. CBS Biodiversity Series 1: 457, 2003.
= Phaeoramularia isolonae Siboe, P.M. Kirk & PE Cannon, in Siboe et al.,
Sydowia 52(2): 299, 2000.
ILLUSTRATION: Siboe et al. (2000: 300, Fig. 6).
On Isolona cauliflora, Africa, Kenya.
Passalora miliusae U. Braun & Crous,
in Crous & Braun, Mycosphaerella and its anamorphs. 1. Names published
in Cercospora and Passalora. CBS Biodiversity Series 1: 460, 2003.
= Mycovellosiella indica P. Kumar & Kamal, Curr. Sci. 51: 846, 1982, non Passalora indica
Kamal & P. Kumar, 1981.
ILLUSTRATION: Kumar & Kamal (1982: 846, Figs 1-2).
On Miliusa tomentosa, Asia, India.
Passalora xylopiae (Viégas & Chupp) U. Braun & Crous,
in Crous & Braun, Mycosphaerella and its anamorphs. 1. Names published
in Cercospora and Passalora. CBS Biodiversity Series 1: 432, 2003.
= Cercospora xylopiae Viégas & Chupp, in Viégas, Bol. Brasil. Agron. 8: 58, 1945.
LITERATURE: Chupp (1954: 46).
On Annona dioica and Xylopia grandiflora, South America, Brazil.
Phacellium annonae-cherimoliae, see above.
Pseudocercospora aethiopicae Deighton, Mycol. Pap. 140: 13, 1976.
Illustration: Deighton (1976: 14, Fig. 3).
On Xylopia aethiopica, Africa, Sierra Leone.
218 ... Braun & Crous
Pseudocercospora annonacea (Kamal, Narayan & R.P. Verma) U. Braun,
Nova Hedwigia 58(1-2): 209, 1994.
= Cercoseptoria annonacea [‘anonacea’| Kamal, Narayan & R.P. Verma, Indian Phytopathol.
39(3): 455, 1987 [‘1986’].
= Pseudocercosporella miliusae M.D. Mehrotra & R.K. Verma, Mycol. Res. 95: 1168, 1991.
LITERATURE: Braun (1995: 194).
ILLUSTRATION: Kamal et al. (1987: 455, Fig. 2).
On Miliusa velutina, Asia, India.
Pseudocercospora annonae U. Braun & Crous, in Braun et al.,
Feddes Repert. 113: 117, 2002.
= Cercospora annonae [‘anonae’| A.S. Mull. & Chupp, Arq. Inst. Biol. Veg. Rio de Janeiro 1:
214, 1935, nom. inval.
LITERATURE: Chupp (1954: 45), Crous & Braun (2003: 59).
ILLUSTRATION: Braun et al. (2002: 115, Fig. 5).
On Annona cherimolia, A. muricata, A. reticulata, A. squamosa, Annona sp.,
Asia, West Indies, Central to South America (Brazil, Cuba, Guatemala, India,
Panama, Philippines, Venezuela).
Pseudocercospora annonae-squamosae U. Braun & R.F. Castafieda,
in Castafieda & Braun, Cryptog. Bot. 1: 50, 1989. Fig. 4
= Cercospora caracasensis Chupp & A.S. Mull., Bol. Soc. Venez. Ci. Nat. 8: 39, 1942, nom.
inval.
= Pseudocercospora annonicola Goh & W.H. Hsieh, in Hsieh & Goh, Cercospora and similar
fungi from Taiwan: 22, 1990.
LITERATURE: Chupp (1954: 45), Vasudeva (1963: 67).
ILLUSTRATIONS: Vasudeva (1963: 68, Fig. 33), Castafeda & Braun (1989: 48, Fig. 19),
Hsieh & Goh (1990: 22, Fig. 10).
MATERIAL EXAMINED: CUBA. Santiago de las Vegas, on living leaves of Annona
squamosa, 29 Dec. 1987, R.E. Castafieda C87/376 (HAL 1650 F), holotype of P annonae-
squamosae. BRAZIL. State of Ceara, Paraipaba City, on living leaves of Annona cherimolia
x squamosa, 19 May 2003, E. Cordoso (HAL 2219 F); State of Ceara, Pacajus County,
on living leaves of Annona muricata, 19 Sep. 1999, F. Freire (HAL 2222 F); State of Para,
Amazonian Region, Belém City, on living leaves of Annona pygmaea, 30 Aug. 2003, L.
Poltronieri (HAL 2220 F); State of Ceara, Pacajus County, on living leaves of Annona
squamosa, 11 May 1999, F. Freire (HAL 2223 F); State of Ceara, Cascavel County, Preaoca
District, on living leaves of Annona sp., 16 Mar. 2000, F. Freire (HAL 2224 F); State
of Para, Amazonian Region, Belém City, on living leaves of Rollinia mucosa, 28 Aug.
2003, L. Poltonieri (HAL 2221 F). VENEZUELA. Caracas, on living leaves of Annona
purpurea, 1 Mar. 1938, A.S. Muller (CUP-VZ 2111), holotype of C. caracasensis.
On Annona atemoya, A. cherimolia x squamosa, A. muricata, A. purpurea,
A. pygmaea, A. reticulata, A. squamosa, Annona sp., Rollinia mucosa, Asia, West
Indies, central to South America (Brazil, Cuba, India, Japan, Panama, Taiwan,
Venezuela).
Cercosporoids on Annonaceae ... 219
Fig. 4. Pseudocercospora annonae-squamosae. A. Conidiophore fascicles. B. Superficial hyphae with
solitary conidiophores. C. Hyphae climbing a leaf hair. D. Conidia (A based CUP-VZ 2111, B-D
based on type material. Scale bar = 10 um. U. Braun del.
ComMENts: With regard to the formation of solitary and fasciculate
conidiophores, P. annonae-squamosae is rather polymorphous. Relatively large,
dense, sometimes almost sporodochial fascicles of conidiophores with well-
220 ... Braun & Crous
developed stromata, are usually epiphyllously formed, whereas on the lower
leaf side the conidiophores are usually solitary arising from superficial hyphae,
and only arranged in smaller fascicles with lacking or small stromata. The
development of superficial hyphae with solitary conidiophores on the one hand
and well-developed fascicles on the other hand is rather variable. Superficial
mycelium may be sparsely developed in some collections, as for instance in
the type material of C. caracasensis, and well-developed conidiophore fascicles
may be lacking in other specimens, as in the holotype of this species. Therefore,
the following redescription of this species based on a wider range of collections
on various host species is necessary in order to demonstrate its variability:
LEAF Spots amphigenous, subcircular to angular-irregular, 1-30 mm diam.,
occasionally confluent, reddish brown, medium to dark or blackish brown,
later often grayish brown to dingy gray, finally sometimes grayish white, often
vein-limited, margin indefinite or with a darker marginal line. CAESPITULI
amphigenous, on the upper side conspicuous, punctiform, scattered, dark
brown to blackish, but soon turning grayish by abundant conidial formation,
on the lower leaf surface subeffuse and less conspicuous, dingy grayish
olivaceous to olivaceous-brown. MyYcELIuM internal and external; superficial —
hyphae variable, ranging from almost lacking to well-developed, usually
hypophyllous, emerging through stomata, occasionally climbing leaf hairs,
branched, septate, 1-3.5 um wide, thin-walled, pale olivaceous to olivaceous-
brown, smooth. Stromata almost lacking to well-developed, above all on the
upper side, 10-80 um diam., substomatal to intraepidermal, olivaceous-brown.
CONIDIOPHORES in small, loose to large and dense, almost. sporodochial
fascicles, arising from internal hyhae or stromata, emerging through stomata
or erumpent, and solitary, arising from superficial hyphae, lateral, rarely
terminal (on the upper side usually in well-developed, sometimes almost
sporodochial fascicles, on the lower side in small, loose fascicles and solitary,
arising from superficial hyphae), erect, straight, subcylindrical, conical to
distinctly geniculate-sinuous, usually unbranched, 5--30(-45) x 2-4(-5) um,
0-1(-2)-septate, subhyaline to pale olivaceous or olivaceous-brown, medium
olivaceous-brown in mass, thin-walled, smooth. CONIDIOGENOUS CELLS
integrated, terminal or conidiophores reduced to conidiogenous cells, 5-20 um
long; conidiogenous loci inconspicuous to subconspicuous by being truncate
or subdenticulate, but always unthickened and not darkened. Conrp1a solitary,
narrowly obclavate-cylindrical, (15—)20-75(-85) x (1.5—)2—4(-5) um, (1-)2-6
(—8)-septate, occasionally slightly constricted at the septa, subhyaline to pale
olivaceous, thin-walled, smooth, apex obtuse, above all in cylindrical conidia,
to subacute, above all in obclavate conidia, base short obconically truncate, 1-2
um wide, hila unthickened, neither darkened nor refractive.
Pseudocercospora annonarum, see above.
Cercosporoids on Annonaceae ... 221
Pseudocercospora annonifolii (Bat. & Peres) U. Braun & O.E Freire,
in Braun, Bibl. Lichenol. 86: 82, 2003.
= Cercospora annonifolii Bat. & Peres, Anais do XIII Congresso da Sociedad Botanica do
Brasil: 374, 1964 [*1962’].
Illustration: Braun (2003: 93, Fig. 4).
On Annona crassifolia, South America, Brazil.
Pseudocercospora asiminae (Ellis & Morgan) U. Braun & Crous, comb. nov.
MycoBank, MB 511860.
Bas: Phloeospora |*Phleospora’| asiminae Ellis & Morgan, in Martin, J. Mycol. 3: 88, 1887.
= Rhopaloconidium asiminae (Ellis & Morgan) Petr., Sydowia 6: 301, 1952.
= Miuraea asiminae (Ellis & Morgan) Arx & Constant., in Arx, Proc. Kon. Ned. Akad.
Wetensch., C 86: 39, 1983.
= Cercospora asiminae Ellis & Kellerm., J. Mycol. 3: 103, 1887.
= Centrospora asiminae (Ellis & Kellerm.) Deighton, Mycol. Pap. 124: 5, 1971.
= Mycocentrospora asiminae (Ellis & Kellerm.) Deighton, Taxon 21: 716, 1972.
Literature: Chupp (1954: 45), Braun (1995: 223), Crous & Braun (2003: 68).
Illustration: Braun (1995: 222, Fig. 211a).
On Asimina parviflora, ‘A. obtusa and A. triloba, North America, USA.
COMMENTS: Based on a re-examination of type material of Phloeospora
asiminae and Cercospora asiminae, Braun (1995) confirmed that the two taxa
are conspecific and, due to the occurrence of some conidia with oblique or
longitudinal septa, followed Arx & Constantinescu (in Arx 1983) in placing
this species in Miuraea Hara. The latter genus is insufficiently known and
undoubtedly heterogeneous (Braun 1995). Cultures and molecular sequence
analyses of M. degenerans (Syd. & P. Syd.) Hara, the type species, are not yet
available. This species is characterized by having pigmented, thick-walled,
dictyosporous conidia. Miuraea persicae (Sacc.) Haraisacolorless Mycosphaerella
anamorph probably closely related to the genus Pseudocercosporella Deighton.
The fungus on Asimina species is, however, better placed in Pseudocercospora.
The conidiophores are formed singly, arising from superficial hyphae, as well
as fasciculate, pigmented, with unthickened, neither darkened nor refractive
conidiogenous loci. The conidia are only slightly thickened, transversely septate,
only occasionally with 1(-2) oblique or longitudinal septa (Braun 1995). The
formation of oblique or longitudinal septa in conidia of Pseudocercospora is
unusual, but may occur, even in P. vitis (Lév.) Speg. (Deighton 1976), the type
species, i.e. this feature does not exclude this cercosporoid Asimina species
from the latter genus.
Pseudocercospora asiminae-pygmaeae U. Braun, A monograph of Cercosporella,
Ramularia and allied genera (phytopathogenic hyphomycetes), Vol. 1: 223, 1995.
Illustration: Braun (1995: 222, Fig. 211b).
On Asimina angustifolia, A. incana and A. pygmaea, North America, USA.
222 ... Braun & Crous
Pseudocercospora oblecta (Syd.) Crous & U. Braun, comb. nov.
MyYcCoBANK MB 511861.
Bas.: Cercospora oblecta Syd., Ann. Mycol. 33: 235, 1935.
= Stigmina oblecta (Syd.) Crous & U. Braun, Mycotaxon 57: 285, 1996.
Literature: Chupp (1954: 45).
Illustration: Crous & Braun (1996: 286, Fig. 8).
On Annona senegalensis, South Africa.
COMMENTSs: On account of rather coarse conidiophores with percurrently
proliferating conidiogenous cells, Crous & Braun (1996) assigned -C. oblecta
to the genus Stigmina Sacc. in the broad sense of Ellis (1971, 1976). The
conidiogenous loci in this species are pseudocercospora-like, i.e. unthickened
and non-pigmented. Braun (1993) introduced the genus Cercostigmina
U. Braun for stigmina-like Mycosphaerella anamorphs with thin-walled,
euseptate, scolecosporous conidia, which was later reduced to synonymy with
Pseudocercospora based on a molecular and morphological reassessment (Crous
& Braun 2003). Scolecostigmina U. Braun (Braun et al. 1999) was proposed for
similar anamorphs with coarse, thick- and often rough-walled conidiophores
formed in sporodochial conidiomata and relatively broad conidia, with slightly
thickened, relatively dark conidia. The status of this genus is unclear. Cultures
of the type species and molecular sequence analyses are not yet available. But
most species assigned to this genus are probably Mycosphaerella anamorphs
that should better be accommodated in Pseudocercospora. Crous et al. (2006)
demonstrated that even Stigmina platani (Fuckel) Sacc., the type species of
Stigmina, characterized by having phragmosporous, distoseptate conidia,
represents a Mycosphaerella anamorph clustering in the Pseudocercospora
clade. Braun & Crous (2006) proposed to conserve Pseudocercospora against
Stigmina and Phaeoisariopsis. Cercospora oblecta is morphologically similar to
Pseudocercospora annonifolii.
Pseudocercospora polyalthiae J.M. Yen, A.K. Kar & B.K. Das,
Mycotaxon 16(1): 43, 1982.
Illustration: Yen et al. (1982: 44, Fig. 4 E-G).
On Polyalthia suberosa, Asia, India.
Pseudocercospora xenoannonicola Crous & Bench.,
in Crous et al., Sydowia 52(2): 88, 2000.
Illustration: Crous et al. (2000: 87, Fig. 10).
On Annona montana, South America, Brazil.
Stenella annonaceae, see above.
Cercosporoids on Annonaceae ... 223
Stenella polyalthiae [‘polylathiae’] S. Chaudhary, N. Sharma & Kamal,
Indian Phytopathol. 55(1): 58, 2002.
Illustration: Chaudhary et al. (2002: 58, Fig. 2).
On Polyalthia suberosa, Asia, India.
Acknowledgements
Thanks are due to the curators of the herbaria CUP and W for the possibility to
examine type material in their keeping. We are much obliged to Chiharu Nakashima,
Mie University, Japan, and Roland Kirschner, J. W. Goethe University Frankfurt a. M.,
Germany, for providing presubmission reviews.
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Braun U. 1993. New genera of phytopathogenic deuteromycetes. Cryptogamic Botany 4: 107-114.
Braun U. 1995. A monograph of Cercosporella, Ramularia and allied genera (phytopathogenic
hyphomycetes). Volume 1. Eching.
Braun U. 1998. A monograph of Cercosporella, Ramularia and allied genera (phytopathogenic
hyphomycetes). Volume 2. Eching.
Braun U. 2003. Miscellaneous notes on some cercosporoid hyphomycetes. Bibliotheca Lichenologica
86: 79-98.
Braun U, Crous PW. 2006. (1732) Proposal to conserve the name Pseudocercospora against Stigmina
and Phaeoisariopsis (Hyphomycetes). Taxon 55(3): 803.
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of All-Russian Institute for Plant Protection (LEP). Mikologia i Fitopatologia 30(4): 1-9.
Braun U, Mouchacca J, McKenzie EHC. 1999. Cercosporoid hyphomycetes from New Caledonia
and some other South Pacific islands. New Zealand Journal of Botany 37: 297-27.
Braun U, Crous PW, Pons N. 2002. Annotated list of Cercospora species (epithets a-b) described by
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1; 42-55.
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57-60.
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author.
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and Passalora. CBS Biodiversity Series 1: 1-571.
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Mycosphaerella from South America. Sydowia 52(2): 78-91.
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Crous PW, Liebenberg MM, Braun U, Groenewald JZ. 2006. Re-evaluating the taxonomic status
of Phaeoisariopsis griseola, the causal agent of angular leaf spot of bean. Studies in Mycology
55: 163-174.
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Cif. and Cercoseptoria Petr. Mycological Papers 140: 1-168.
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Hennings P. 1908 [*1909’]. Fungi S. Paulenses IV. Hedwigia 48: 1-20.
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World. 8th edn. Regnum vegetabile 120: 1-163.
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Phytopathology 39(3): 453-458.
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846-847.
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Siboe GM, Kirk PM, David JC, Cannon PE. 2000. Necrotrophic fungi from Kenyan endemic and
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Yen JM, Kar AK, Das BK. 1982. Studies on Hyphomycetes from West Bengal, India, I. Cercospora
and allied genera of West Bengal, 1. Mycotaxon 16(1): 35-57.
MYCOTAXON
Volume 105, pp. 225-234 July-September 2008
Novelties on Southern Brazilian Parmeliaceae
MARCELO P. MARCELLI & LUCIANA S. CANEZ
mpmarcelli@msn.com & lucanez@yahoo.com.br
Instituto de Botanica, Se¢do de Micologia e Liquenologia,
Caixa Postal 3005, Sao Paulo / SP 01061-970, Brazil
Abstract — Bulbothrix megapotamica, Myelochroa supraflava and Xanthoparmelia
conglomerata are described as new species. The new combination Parmotrema aberrans
is made. Rimelia luminosa is synonymized under Parmotrema diffractaicum.
Key words — Bulbothrix subcoronata, Myelochroa aurulenta, Xanthoparmelia
punctulata
The Parmeliaceae is the most significant family of lichenized fungi in the
Brazilian biomes (Marcelli 1998). The same is true for Rio Grande do Sul State,
where 169 species in 17 genera were already reported (Spielmann 2006).
During a survey of the Parmeliaceae in the area of Fazenda da Estrela (Canéz
2005) several new species were found and three of them are formally described
here. In addition, a new combination and a new synonym are proposed.
Fazenda da Estrela, with an area of 287 km’, is located in Vacaria Municipality,
in the northern highlands of the southern Brazilian state of Rio Grande do
Sul. Araucaria angustifolia (Bertol.) Kuntze forests and grasslands are the
predominant vegetation types.
The material and methods used here are described in Canéz & Marcelli
(2007).
New species
Bulbothrix megapotamica Canéz & Marcelli, sp. nov. Fic. 1
MycoBank MB 511099
Diacnosis: Sicut Bulbothrix subcoronata, sed sporis majoribus differt.
Ho otypre-Brazil, Rio Grande do Sul State, Municipality of Vacaria, Fazenda da Estrela,
28°04’58.9”S, 50°56’48.9” W, 850 m alt., open field, on a shrub branch, leg. L.S. Canéz &
A.A. Spielmann 146, 23-I-2003 (SP).
THALLUS whitish gray, sublaciniate, up to 5.0 cm wide; SUBLACINIAE irregularly
branched, 0.4-1.5 mm wide, adnate, contiguous, apices slightly truncate, often
226 ... Marcelli & Canéz
recurved, almost perpendicular to the substrate, margin smooth, surface
continuous, smooth, slightly shiny. LACINULAE and MACULAE absent. CILIA
black, simple, 0.15-0.35 x 0.05 mm, bulbate, bulbs 0.05-0.20 mm wide,
abundant, contiguous on the axils, with interciliary spaces equivalent to one
bulb on the lateral margins and almost contiguous at the apices. PUSTULAE,
SORALIA and IsIDIA absent. MEDULLA white throughout. LOWER SURFACE
black, shiny, slightly rugose; MARGINAL zone concolorous or dark brown, shiny,
0.45-0.75 mm wide, poorly delimited, smooth; RHIzINAE black, present to the
margin, simple, base not bulbate, 0.20-0.75 x 0.05 mm, very dense and evenly
distributed. APOTHECIA slightly concave, 0.5-3.0 mm diam., adnate, laminal,
margin irregularly crenate and coronate, with almost equally sized bulbae in
the axils, disc brown, slightly concave, imperforate, amphithecium smooth;
ASCOSPORES ellipsoid, 13.7-17.5 x (7.5—) 8.7-10.0 (—11.0) um, epispore 1.2 um
thick. Pycnrp14 laminal and submarginal, ostiole black; conrp1a bifusiform,
4.0-6.0 x 1.0 um.
SPOT TESTS— cortex K+ yellow, UV-; medulla K+ yellow — reddish orange, C-,
KC-, P+ yellow, UV-.
TLC- atranorin (traces, cortex) and norstictic acid (medulla).
CoMMENTS- Bulbothrix megapotamica is distinctive by producing coronate
apothecia with large ascospores (13.7-17.5 um), abundant cilia, the black
lower surface, and the medullary norstictic acid, associated to the absence of
vegetative propagules.
This new species is part of a group of adnate sublaciniate species with globose
cilia and norstictic acid containing, common in Southeast Brazil. It is similar
to B. subcoronata (Mill. Arg.) Hale that has smaller ascospores (5-7.5 um),
and B. viatica Spielmann & Marcelli with ascospores 12-17.5 um but a brown
undersurface (Spielmann & Marcelli 2008).
Miller Argoviensis reported the ascospores of B. subcoronata as 5 um long,
the smallest size found in the genus. Several authors apparently considered that
a mistake, as Jungbluth (2006) reported sizes from 8 to 15 um based on reports
by Eliasaro (2001), Fleig (1985) and Marcelli (1993) for material identified as B.
subcoronata from several places in natural areas of southern and southeastern
Brazil. However, a recent examination of the lectotype (G!) of B. subcoronata
revealed that the spores are really 5-7.5 um long (Benatti, pers. comm.).
Furthermore, influenced by Hale's classical monograph on Bulbothrix, .
all these researchers have named as B. subcoronata those specimens with a
completely brown underside. However, the lectotype analysis revealed that the
real underside color is of an undeniable bright black, with only a narrow brown
marginal zone.
New species of Parmeliaceae (Brazil) ... 227
The concept of B. subcoronata is currently under revision by M.N. Benatti,
and we believe that at least two or three species with different ascospore size
and underside color may be involved.
Bulbothrix sensibilis (J.Steiner & Zahlbr.) Hale, a species with black underside,
has cilia with short apices and salazinic acid in the medulla. All other similar
species have smaller ascospores and/or a brownish underside.
The holotype of B. megapotamica has slightly downturned cilia almost
perpendicular to the substrate; however, it was not possible to decide if this is a
character of taxonomical value.
The specific epithet (Greek: mega + potamos = big river), several times used
by G.O. Malme for species of the same region, refers to Rio Grande do Sul State,
whose name means Big South River.
Myelochroa supraflava Canéz & Marcelli, sp. nov. Fig. 2
MycoBAnk MB 511098
Diacnosis: Affinis Myelochroae aurulentae, sed medulla supra lutea et infra alba differt.
Ho.otyPe-Brazil, Rio Grande do Sul State, Municipality of Vacaria, Fazenda da Estrela,
riverside vegetation, 28°04’49.2”S, 50°56'36.9”W, 850 m alt., saxicolous (basalt), near
the stream, leg. L.S. Canéz & A.A. Spielmann 1377, 10-I-2004 (SP; isotypes in HAS and
NY).
THALLUS whitish gray, sublaciniate, up to 9 cm in diameter; LACINIAE
subdichotomous, 0.5-3.0 mm wide, adnate, contiguous to slightly overlapping,
apices brownish, mostly truncate, margin smooth, upper surface continuous,
smooth but slightly irregularly pitted, shiny; LACINULAE absent; MACULAE
weak, punctiform, laminal; c1i1A black, simple to rarely furcate, 0.10-0.25 x
0.05 mm, irregularly dispersed throughout at the margin but contiguous in the
axillae. PUSTULAE small, generally 0.1-0.3 mm, mostly subapical, sometimes
developed also on elevated parts of the lamina, circular to slightly elongated,
bursting apically since young to form coarse soredia and often leaving the
yellow medulla exposed, chiefly aggregated into capitate masses 1.0-2.5 mm
diam. SORALIA capitate, originating from the pustulae; soREDIA granular and
subgranular. Isrp1A absent. MEDULLA bicolored, the upper 3/4 pigmented
yellow and the lower 1/4 portion white. Lower surFACE black, shiny, rugose;
MARGINAL ZONE dark brown, shiny, 0.3-1.5 mm, poorly delimited, densely
papillate; RHIZINAE black, simple to less frequently irregularly branched,
0.25-0.90 mm long, dense, almost reaching the margins, often with grouped
to agglutinated, especially in the thallus center. APOTHECIA absent. PYCNIDIA
absent.
SPOT TESTS— cortex K+ yellow, UV- or + pale yellow on the tips of some lobes;
medulla K+ burnt yellow, C+ strongly yellow, KC-, P-, UV-.
228 ... Marcelli & Canéz
TLC-— atranorin (cortex), zeorin, secalonic acid A and leucotylic acid
(medulla).
Paratypes: Brazil, Rio Grande do Sul State, Municipality of Boqueirao do Leao, open
place, 29°18°13.7"S, 52°26'51.7”W, 500 m alt., saxicolous at the margin of a stream,
partially open, leg. A.A. Spielmann & M.A. Sulzbacher 760, 31-I-2004 (SP); idem, Minas
Gerais State, Municipality of Ouro Preto, northeastern outskirts of city Ouro Preto,
20°23’S, 43°29-30’W, 1250 m alt., on hardpan, leg. G. Eiten 6932, 28-IX-1965 (SP).
CoMMENTS- Myelochroa supraflava has oval axils that give it a laced aspect,
resembling some growth patterns of Hypotrachyna livida (Taylor) Hale. The
principal characteristic is its medullary coloration, which is almost completely
yellow, but with a thin white layer adjacent to the lower cortex. The specific
epithet refers to this.
The closest species is M. aurulenta (Tuck.) Elix & Hale, which has the same
chemistry and is sorediate. Compared to M. supraflava, ithasa darker and grayish
green thallus, overlapping lobes, a distinctly maculate surface, some pruinose
lobe apices and a chiefly white medulla, with a yellow or orange color present
only below or around the soralia (Hale 1976b, Elix 1994, Divakar & Upreti
2005). According to Hale (1976b), M. aurulenta has sublinear to subirregular
or rounded, 2-4 mm wide lobes, simple to barely forked or squarrose rhizinae,
and distinct, cracked areas that expose the medulla. Myelochroa supraflava also
has these areas; however, it is yellow throughout the upper medulla.
Myelochroa supraflava also resembles the Southeast Asian M. denegans (Nyl.)
Elix & Hale, which differs in having a reddish orange pigment in the lower part
of medulla and pustalae in part entire (Elix 1994, Divakar & Upreti 2005).
Three records of M. aurulenta are published for Brazil (Hale 1976b, Marcelli
1993, Spielmann 2005). The revision of this material demonstrated that
the specimens discussed by Hale and Spielmann are M. supraflava, and that
Marcelli’s specimen is a tiny lichen, possibly a new species of Parmelinopsis
with yellow medulla. Apparently, M. aurulenta is not yet known from Brazil.
Xanthoparmelia conglomerata Canéz & Marcelli, sp. nov. Fic. 3
MycoBank MB 511100
Diaenosis: Species thallo ut in Xanthoparmelia punctulata sed ab hac specie superficie
inferiore nigra, isidiis globularibus deinde agglomeratis et acidum salazinicum continenti
differt.
Ho.otype-Brazil, Rio Grande do Sul state, Municipality of Vacaria, Fazenda da Estrela,
field, 28°02’44.6"S, 51°02’01.7”W, 860 m alt., on basaltic rock in a sunny place at the
Frade River margin, leg. L.S. Canéz & A.A. Spielmann 692a, 11-I-2004 (SP; isotype in
B).
THALLUS greenish yellow, laciniate, 8.0-13.0 cm wide; LACINIAE subdi-
chotomically or irregularly branched, 0.7-3.0 mm wide, separate to laterally
imbricate, apices round to truncate, darkened; margins smooth; upper surface
New species of Parmeliaceae (Brazil) ... 229
Fig. 1. Bulbothrix megapotamica (part of the holotype).
Fig. 2. Myelochroa supraflava (part of the holotype).
Fig. 3. Xanthoparmelia conglomerata (part of the holotype).
Bars = 5 mm.
230 ... Marcelli & Canéz
continuous, rarely cracked in the central region, mostly smooth, shiny on the
younger parts. LACINULAE, MACULAE, PUSTULAE and SORALIA absent. IsipDIA
concolorous with the thallus, sometimes with brownish apices, mostly globose
when young, then proliferating into large, capitate, sometimes coralloid,
aggregates, 0.10—0.35 x 0.10-0.25 mm, erect, firm, eciliate at apices, laminal,
rarely marginal. MEDULLA white, pigments absent. LOwER suRFACE black to
dark brown, opaque, smooth; MARGINAL ZONE dark brown, shiny, 2.0—5.0(—7.0)
mm wide, margin poorly delimited, smooth to slightly rugose; RHIZINAE black
or concolorous with the margin, simple, 0.25-0.75 x 0.05-0.15 mm, unevenly
distributed. APOTHECIA concave to almost plane, 0.75-1.50 mm in diameter,
adnate, laminal, margins smooth to crenate, amphithecium smooth or barely
isidiate, disc dark brown, imperforate; AscosPoREs absent. PycNnip14 laminal
and submarginal, ostiole black; conrp1a bifusiform, 5.0-7.5 x 1.0 um.
SPOT TESTS— cortex K-, UV-; medulla K+ yellow — red, C-, KC-, P+ yellow,
UV- or + pale yellow.
TLC-— usnic acid (cortex), salazinic acid and traces of consalazinic acid
(medulla).
PARATYPE—Brazil, Rio Grande do Sul State, Municipality of Vacaria, Fazenda da
Estrela, field, 28°02°44.6"S, 51°02’01.7" W, 860 m alt., on basalt in sunny area at
the Frade River margin, leg. L.S. Canéz & A.A. Spielmann 692b, 1 1-1-2004 (SP).
CoMMENTS-— Xanthoparmelia conglomerata is characterized by the aggregates
of globular isidia, the production of salazinic acid (K+ yellow — red) and the
black undersurface.
The species is easily recognized by its globose isidia that proliferate into
large aggregates, which have a typical popcorn-like aspect (the reason for the
name). Sometimes the younger isidia may look like granular soredia; however,
an accurate examination reveals that they are truly granular isidia, that are
completely corticate.
Xanthoparmelia tinctina (Maheu & A. Gillet) Hale, also with black
undersuface and medullary salazinic acid, has different barrel-shaped to
irregularly inflated isidia and a lobate thallus; in addition, their isidia do not
form agglomerates and produces additionally norstictic acid (Hale 1990).
Xanthoparmelia antleriformis (Elix) Elix & J. Johnst. differs by its dark brown
lower surface, its ramified, coralloid to antleriform isidia, and the presence of
traces of norstictic acid together with salazinic and consalazinic acid. (Nash
et al. 1995). Xanthoparmelia punctulata (Gyeln.) Hale produces salazinic and
consalazinic acids in the medulla and has globose isidia; however, they are
erumpent and form scurfy masses (Hale 1990). Xanthoparmelia glomerulata
Krog & Swinscow has a brown lower surface and obconical isidia that degrade
to corticated granules, and produce the stictic acid complex (Krog & Swinscow
1987).
New species of Parmeliaceae (Brazil) ... 231
New combination
Parmotrema aberrans (Vain.) Canéz & Marcelli, comb. nov.
MycoBank MB 511103
BasIonyM: Parmelia xanthina f. aberrans Vain., Acta Societatis pro Fauna et Flora
Fennica 7(1): 37. 1890.
Type—Brazil, Minas Gerais State, Sitio [nowadays Anténio Carlos Municipality], leg.
E.A. Vainio s.n., Lichenes brasiliensis exsiccati 664 (TUR-V 02758, holotype).
COMMENTS-—Parmotrema aberrans is characterized by a greenish yellow (usnic
acid in the cortex) thallus, continuous cilia along the margins, cylindrical, ciliate
isidia, and by producing gyrophoric acid in the medulla (C+ and KC+ rose).
Parmotrema xanthinum (Mill. Arg.) Hale also has cortical usnic acid and
cylindrical, ciliate isidia, but does not produce gyrophoric acid (medulla C-,
KC-) (Hale 1965, Nash & Elix 2002).
In his classical monograph, Hale (1965) mentioned Parmelia xanthina
(Mull. Arg.) Vain. as having no medullary reactions, and placed Parmelia
(= Parmotrema) madagascariacea (Hue) Abbayes (medulla C- and KC+ rose)
under it as a synonym, distinguishing Parmelia aberrans (Vain.) Abbayes by its
positive medullary reactions.
In order to understand the chemistry and nomenclature of the P xanthina
complex, Almeda & Dey (1973) made chromatography and agreed with Hale
(1965) in naming P. xanthina the group without medullary reactions. They
also discovered that the C-, KC+ rose collections in fact produced gyrophoric
acid and had a subtle C+ rose reaction that could be easily overlooked. For
this reason, they considered ‘Parmelia aberrans Hale’ a synonym of Parmelia
madagascariacea, the oldest valid combination.
Some years later, Krog & Swinscow (1981) made chromatography of the
Parmelia madagascariacea holotype and mentioned the presence of an “acid
aff. gyrophoric” (gyrophoric complex) different from gyrophoric acid in the
holotype of P. aberrans. However, they considered both P aberrans and Parmelia
madagascariacea as synonyms of P. xanthina, not attributing any taxonomic
value to the presence of gyrophoric acid.
Parmotrema aberrans was reported for Rio Grande do Sul State by Fleig
(1990), from the Esmeralda Municipality. Years later, she (Fleig 1997) considered
the species as synonym of P. xanthinum, and credited the combination
Parmotrema aberrans to Hale (mentioning Hale 1965). However, Hale (1974)
recognized the genus Parmotrema only about 10 years later and never made
this combination.
Recently, Nash & Elix (2002) recognized and cited the species as ‘Parmotrema
aberrans (Vain.) Abbayes’ for the Sonoran Desert, but Abbayes made the
combination Parmelia aberrans and never attributed the taxon to the genus
Parmotrema (Abbayes 1959, according to Hale 1965).
232 ... Marcelli & Canéz
Specimens of Parmotrema aberrans cited as P. xanthinum are also known from
the mountains of Sao Paulo and Minas Gerais States in southeastern Brazil
(Ribeiro 1998). )
Recently, Canéz (2005) and Jungbluth (2006) recognized Parmotrema
aberrans, from Rio Grande do Sul Araucaria forests and Sao Paulo State cerrado
vegetation, respectively, but did not publish a valid combination.
As a matter of fact, there is no reliable report of the real P xanthinum from
Brazil, perhaps not even from South America.
New synonym
Parmotrema diffractaicum (Essl.) Hale, Phytologia 28(4): 335. 1974.
BastonyYM: Parmelia diffractaica Essl., Bryologist 75(1): 80. 1972.
SyNONYM-—Rimelia luminosa Marcelli, Mitteilungen aus dem Institut fiir Allgemeine
Botanik Hamburg 30-31: 152. 2002. [Holotype: Brasil, Minas Gerais, municipio
de Catas Altas, Serra do Caraca, Parque Natural do Caraga, trilha para Cascatona,
sobre tronco de arvore na mata umida, 1330 m alt., 11-I[X-1997, col. Marcelli &
Ribeiro 32020 (SP !, holotype; B, isotype)].
COMMENTS — Parmotrema diffractaicum is characterized by soralia and
diffractaic acid associated with lichexanthone in the medulla (UV+ yellow
orange-colored).
Morphologically, it is somewhat similar to Parmotrema spinibarbe (Kurok.)
Hale ex DePriest & B.W. Hale, which also has medullary lichexanthone but
produces additionally salazinic acid and its soralia is frequently submarginal,
subapical or laminal.
Rimelia luminosa in the original description by Marcelli & Ribeiro (2002)
was stated to differ in forming soralia and producing lichexanthone associated
with a-collatolic acid in the medulla. However, chromatography of the
holotype demonstrated medullary diffractaic acid instead, and its morphology
is the same as described for P. diffractaicum. Consequently, we consider both
as synonyms.
Acknowledgements
The authors wish to thank Ana Crespo (Madrid), Thomas H. Nash (Tempe), and
Harrie J. M. Sipman (Berlin) for the critical revision of the manuscript, and Teuvo
Ahti (Helsinki) for the revision of Latin diagnoses. We are also thankful to Adriano A.
Spielmann for the photo of B. megapotamica and Michel Navarro Benatti for provide
original data of his thesis.
We acknowledge the Conselho de Desenvolvimento Cientifico e Tecnolégico (CNPq)
for a research grant to the first author and the Coordenacao de Aperfeicoamento de
Pessoal de Nivel Upper (CAPES) for supporting a Master's Scholarship for the second
author.
New species of Parmeliaceae (Brazil) ... 233
Literature cited
Almeda F Jr, Dey JP. 1973. Chemical and nomenclatural notes on the Parmelia xanthina complex.
Bryologist 76: 541-543.
Canéz LS. 2005. A familia Parmeliaceae na localidade de Fazenda da Estrela, municipio de Vacaria,
Rio Grande do Sul, Brasil. Mastership dissertation, Instituto de Botanica, Sao Paulo, 302 p.
Canéz LS, Marcelli MP. 2007. Two new species of Punctelia (Parmeliaceae) from southern Brazil.
Mycotaxon 99: 211-216.
Divakar PK, Upreti DK. 2005. Parmelioid lichens in India. A revisionary study. India: Bishen Singh
Mahendra Pal Singh, 488 p.
Eliasaro S. 2001. Estudio taxonomico y floristico sobre las Parmeliaceae sensu stricto (Ascomycota
Liquenizados) del Segundo Planalto del Estado de Parana, Brasil. Buenos Aires. Doctoral
thesis (en Ciencias Bioldgicas). Universidad de Buenos Aires, Facultad de Ciencias Exactas y
Naturales.
Elix JA. 1994. Myelochroa. Flora of Australia 55: 66-67.
Fleig M. 1985. Estudo preliminar da familia Parmeliaceae (liquens) no Rio Grande do Sul, Brasil.
Comunica¢des do Museu de Ciencias e Tecnologia da PUCRS, Série Botanica, 35: 79-91.
Fleig M. 1990. Liquens da Estacao Ecoldégica de Aracuri. Novas ocorréncias no Rio Grande do Sul.
Theringia Série Botanica 40: 121-125.
Fleig M. 1997. Os géneros Parmotrema, Rimelia e Rimeliella (Lichenes - Ascomycotina,
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Biociéncias, Universidade de Sao Paulo.
Hale ME. 1965. A monograph of Parmelia subgenus Amphigymnia. Contributions from the United
States National Herbarium 36: 193-358.
Hale ME. 1974. New combinations in the lichen genus Parmotrema Massalongo. Phytologia 28:
334-339.
Hale ME. 1976a. A monograph of the lichen genus Bulbothrix Hale (Parmeliaceae). Smithsonian
Contributions to Botany 32: 1-29.
Hale ME. 1976b. A monograph of the lichen genus Parmelina Hale (Parmeliaceae). Smithsonian
Contributions to Botany 33: 1-60.
Hale ME. 1990. A synopsis of the lichen genus Xanthoparmelia (Vainio) Hale (Ascomycotina,
Parmeliaceae). Smithsonian Contributions to Botany 74: 1-250.
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102: 462-544.
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Estado de Sao Paulo. Mastership dissertation. Instituto de Botanica. Sao Paulo. 323 p.
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of the British Museum, Botany Series 9: 143-231.
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genera, with special emphasis on East African taxa. Lichenologist 19: 419-431.
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Acta botanica brasilica 7: 25-70.
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MRD. (eds). Lichenology in Latin America: history, current knowledge and applications, Sao
Paulo: CETESB, p. 25-45.
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125-155:
234 ... Marcelli & Canéz
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Lichen Flora of the greater Sonoran Desert Region. Vol. 1. Lichens Unlimited, Arizona State
University, Tempe, Arizona, USA. pp. 318-329.
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America. Bibliotheca Lichenologica 56: 1-157.
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dos Estados de Minas Gerais, Rio de Janeiro e Sao Paulo. Mastership dissertation. Instituto de
Biociéncias, Universidade de Sao Paulo, 194 p.
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encosta da Serra Geral, no Vale do Rio Pardo, Rio Grande do Sul, Brasil. Mastership dissertation,
Instituto de Botanica, Sao Paulo, 204 p.
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Caderno de Pesquisas Série Biologia 18(2): 7-125.
Spielmann AA, Marcelli MP. 2008. Bulbothrix viatica, a new species of Parmeliaceae from Brazil.
Mycotaxon 103: 201-205.
MYCOTAXON
Volume 105, pp. 235-248 July-September 2008
New species of Parmotrema containing protocetraric or stictic
acids from the coast of Sao Paulo State, southeastern Brazil
MARCELO P. MARCELLI & MICHEL N. BENATTI
mpmarcelli@msn.com
Instituto de Botanica, Secao de Micologia e Liquenologia
Caixa Postal 3005, Sao Paulo / SP 01061-970, Brazil
JOHN A. ELIx
John.Elix@anu.edu.au
Department of Chemistry, Building 33, Australian National University
Canberra, A.C.T. 0200, Australia
Abstract — During a survey of Parmeliaceae species in natural ecosystems and
urbanized coastal areas of southeastern Brazil, three new Parmotrema species were
discovered: P. eliasaroanum, containing stictic acid, and P. graniticum and P. milanezii,
containing protocetraric acid.
Key words — Parmotrema perlatum, Parmotrema dilatatum, Parmotrema gardneri,
Parmotrema lobulatum
Introduction
The genus Parmotrema A. Massal. is characterized by lobes with broad rotund
apices and naked lower margins, the absence of pseudocyphellae, the frequent
occurrence of marginal cilia, simple rhizines, and thick-walled, ellipsoid
ascospores (Brodo et al. 2001, Nash & Elix 2004). More than three hundred
species are known world-wide (Nash & Elix 2004), 93 of which occur in Brazil
(Marcelli 2004).
Two new species containing protocetraric acid and one containing stictic
acid are described here. These species were discovered by the authors during
research on the broad-lobed species of Parmeliaceae at the coast in Sao Paulo
State, Brazil (Benatti 2005), for the most situated between the municipalities
of Ubatuba (23°02’S, 45°04’W) and Itanhaém (24°11’S, 46°47’°W). This region
includes urbanized areas, rocky shores, as well as mangrove and restinga forests
as the predominant vegetation types.
236 ... Marcelli, Benatti & Elix
While Parmotrema graniticum is strictly saxicolous on rocky shores,
the other species are corticolous in coastal mangrove and restinga forests.
Parmotrema eliasaroanum lacks vegetative propagules whereas P. graniticum
and P. milanezii are sorediate.
Although we have included sufficient specific information about the new
taxa, more detailed morphological and chemical comparisons with other
somewhat similar species can be found in Benatti (2005).
Materials and methods
Specimens were distinguished by morphological characters using standard
stereoscopic and light microscopes. Anatomical sections, including those of
apothecia and pycnidia, were made with a razor blade by hand. The chemical
constituents were checked by spot tests with potassium hydroxide (K), sodium
hypochlorite (C) and para-phenylenediamine (P), and also examined under
UV light (360 nm). Chemical constituents were identified by thin-layer
chromatography (TLC) using solvent C (Bungartz 2001), high performance
liquid chromatography (HPLC) (Elix et al. 2003) and comparison with ~
authentic samples.
Since we have encountered problems dealing with the many morphological
terms present in the literature, we specify here that in our concept lacinules
represent adventitious, ribbon-like secondary outgrowths from the primary
lobes margins. Lobules are similar, but short and rounded.
The diagnosis refers exclusively to holotype characters and the English
descriptions and comments to all the material studied.
The species
Parmotrema eliasaroanum Benatti, Marcelli & Elix, sp. nov. Fig. 1
MycoBankK MB 511426
DraGnostis: Species cum thallo simili Parmotrematis lobulati sed thallus magis adpressus
et planus, lobis majoribus et subrotundatis; lacinulis marginalibus planis et ciliatis, non
lobos apicales formantibus; lacinulae magis regulares, non lobulis secundariis; margine
inferiori plerumque eburneo et conidia plerumque minoribus differt. Atranorinam,
chloroatranorinam, acidum sticticum, acidum consticticum, acidum peristicticum, acidum
cryptosticticum et acidum methylsticticum continens.
Ho.otype-Brazil, Sao Paulo State, Municipality of Cananéia, Cardoso Island, restinga
wood of Vila Maruja, 25°14’S, 48°01 ’W, 1 m alt., tree branch at the low wood behind the
dunes, leg. M.P.Marcelli 1757, 20-X-1981 (SP; isotype in B).
THALLUS up to 15 cm in diameter, subcoriaceous, ramulicolous, grayish green
but becoming darker in the herbarium, lobed to sublobed; LoBgs irregularly
branched, (2.0—)3.0-6.0 mm wide, contiguous to + imbricate, becoming
crowded in part when lacinulate, adnate to loosely adnate, lacinules ascending;
Parmotrema spp. nov. (Brazil) ... 237
APICES +flat unless lacinulate, subrotund to irregular; MARGIN smooth or
subcrenate to irregularly dissected, +flat, weakly undulate in part, entire
to incised, lacinulate, ciliate. UPPER SURFACE continuous then weakly and
irregularly cracked, smooth; MACULAE absent; LACINULES present; SORALIA,
PUSTULES and ISIDIA absent; LACINULES short to + elongate, evenly dispersed
at the apices and along the margins of lobes, occasionally becoming crowded,
usually more abundant in older parts, simple to anisotomic or irregularly
dichotomous, + flat, 0.4—5.6 x 0.2-1.2 mm, truncate to subtruncate, ciliate, not
admixed with adventitious lobules, underside cream colored or occasionally
variegated; cia black, simple, 0.2-1.5(—2.3) x ca. 0.05 mm, frequent along
the margins. MEDULLA white, pigments absent. LowER suRFACE black, shiny,
smooth to subrugose, unevenly papillate, erhizinate in parts not in contact
with the substrate; MARGIN shiny to opaque, usually cream or variegated, rarely
pale brown in a few elacinulate distal lobes, smooth to subrugose, 1.0-5.5 mm
wide, naked; RHIZINES black, simple, 0.1-1.3 x ca. 0.05 mm, absent, sparse
or frequent, more common in parts in contact with the substrate, grouped.
APOTHECIA submarginal to subterminal, common, concave, becoming
distorted and involute with age, 0.8-6.0 mm wide, stipitate, margins crenate
to denticulate-lacinulate, eciliate, amphithecia and stipe smooth; pisc brown,
epruinose, imperforate; AscosPorREs ellipsoid, 22.5-29 x 12.5-15 um, epispore
3-4 um wide. Pycnip1A submarginal, common, abundant on lacinules, with
black ostioles; conip1a bacilliform to short filiform, 4-8 x ca. 1 um.
SPOT TESTS: upper cortex K+ yellow, UV-; medulla K+ yellow, C-, KC-, P+
orange, UV-.
TLC/HPLC: cortical atranorin (minor) and chloroatranorin (minor);
medullary stictic acid (major), constictic acid (submajor), peristictic acid
(trace) cryptostictic acid (minor) and methyl stictic acid (minor).
PARATYPE—Brazil, Sao Paulo State, Municipality of Iguape, Barra do Ribeira, between
Suamirim “River” and the ocean, restinga wood at the mangrove’s edge, 24°38’S,
47°22’W, 1 m alt., thin branch of small tree, leg. M.P.Marcelli & O.Yano 6857, 18-07-
1989 (SP).
COMMENTS-Parmotrema eliasaroanum is one of the few lacinulate species
containing the stictic acid chemosyndrome and resembles P. lobulatum Marcelli
& Hale (Fig. 2). However, these species differ in morphology and in some of
their medullary substances.
Parmotrema eliasaroanum is characterized by the adnate thallus with
ascending lacinulate lobes, the ciliate margins, dichotomous or irregularly-
branched, flat, marginal non reproductive lacinules, the absence of soredia and
isidia, an almost entirely cream coloured lower margin (possibly white when
freshly collected), grouped rhizines and the presence of the stictic complex in
238 ... Marcelli, Benatti & Elix
Figures 13.41 Holotype (above) and isotype (below) of P eliasaroanum (a single specimen).
2. Isotype of P lobulatum (SP).
Bar = 1 cm.
Parmotrema spp. nov. (Brazil) ... 239
the medulla. The apothecia are denticulate-lacinulate and eciliate, with smooth
amphithecia.
When initial specimens of P eliasaroanum were examined they were
considered to be P. lobulatum, since the latter has marginal lacinules and stictic
acid as the major medullary substance. However, after studying an isotype (SP!)
and a topotype of this species, consistent differences were observed between
the two species. Parmotrema lobulatum is a smaller lichen with a more loosely
adnate, ascending thallus, with narrower (1.0-3.0 mm vs. 3.0-6.0 mm wide)
and often more crowded lobes. In addition, P lobulatum develops plane, more
irregularly shaped and branched, eciliate lacinules that eventually become
subcanaliculate as well as small, dense, procumbent lobules along the margins
of the lobes and lacinules.
Parmotrema lobulatum also differs from P. eliasaroanum in having very
narrow, brown colored lower margins [even of the lacinulate lobes], its lower
cortex has more scattered rhizines, and the conidia are longer (7-9 um vs. 4-8
um).
The two specimens of P eliasaroanum contain stictic, constictic, peristictic,
cryptostictic and methyl stictic acids, whereas the two specimens of P. lobulatum
contain stictic, constictic, menegazziaic, cryptostictic, peristictic, substictic,
and hypostictic acids.
Other regularly lacinulate species, such as P zicoi Marcelli & C.H. Ribeiro
and P. merrillii (Vain.) Hale, differs in lobe width, lacinule size and shape, the
colour of the lower margins, the presence of cilia, ascospore and conidia size
and by containing protocetraric acid.
This species is named in honor of the Brazilian lichenologist Dr. Sionara
Eliasaro, from the Universidade Federal do Parana, who has made significant
contributions in establishing and developing taxonomic lichenology in Brazil.
Parmotrema graniticum Benatti, Marcelli & Elix, sp. nov. Fig. 3
MycoBank MB 511904
Diacnosis: Species cum thallo simili Parmotrematis mordenii sed thallus magis
coriaceus, cortex superior atranorinam et acidum usnicum continens et medulla acidum
protocetraricum, acidum conprotocetraricum, acidum virensicum, acidum subvirensicum,
et acidum convirensicum continens, sed sine atranorina medullari.
Hototype-Brazil, Sao Paulo State, Municipality of Sao Sebastiao, Centro de Biologia
Marinha da USP (CEBIMAR), Ponta do Baleeiro, southward rocky shore, 23°48’S,
45°37'W, 15 m alt., granitic rock, leg. M.P. Marcelli 2216, 19-ITI-1988 (SP; isotype in
NY).
THALLUS up to 10 cm in diameter, subcoriaceous, saxicolous, greenish gray
becoming darker in the herbarium, lobate to sublobate; Lopes 1.5—-5.0(—7.0)
mm wide, irregularly branched, contiguous to + imbricate, tightly adnate;
APICES +flat, subrotund; MARGIN smooth to crenate or occasionally irregular
240 ... Marcelli, Benatti & Elix
in older parts, becoming more sinuous when sorediate, incised, flat to
+ascending (mainly when sorediate), partially involute, occasionally folded
and subcanaliculate at the axils, t+undulate, unevenly sublacinulate, eciliate.
UPPER SURFACE continuous but becoming irregularly cracked with age, smooth
to subrugose, MACULAE absent, ADVENTICEOUS LACINULES common, very
short and irregularly distributed along the lobes margins, simple, furcate or
irregular, flat, 0.2-1.8(—3.0) x 0.2-0.9 mm, truncate, underside concolourous
with the lower margin, occasionally intermixed with small adventitious
irregular lobules, eciliate; SORALIA mainly marginal and linear, sometimes
also at the apices of the lacinules, occasionally coalescing and becoming
+irregular, sometimes subcapitate and subpustular when (rarely) submarginal
or laminal, sorEpDIA granular to subgranular, not agglutinated. PUSTULEs and
ISIDIA absent. MEDULLA white, pigments absent. LOwER suRFACE black, shiny,
smooth to subrugose, weakly veined in part, MARGIN shiny, reddish brown or
rarely black, smooth, weakly veined, 0.5-4.0 mm wide, naked; RHIzINEs black,
simple, 0.1-0.7 x 0.05-0.1 mm, frequent, occasionally becoming agglutinated,
grouped. APOTHECIA laminal to submarginal, rare, concave, 0.3—2.6 mm wide,
sessile, margins smooth at first then sorediate, eciliate, amphithecia subrugose
to sorediate, stipe smooth to subrugose; pisc brown, epruinose, imperforate;
ASCOSPORES ellipsoid, 7.5-12.5 x 4.-6.5 um, epispore ca. 1 um thick. PycNIDIA
rare, submarginal, with black ostioles; Cconip1a weakly sublageniform,
(4—)5-6.5 x ca. 1 um.
SPOT TESTS: upper cortex K+ yellow, UV-; medulla K- or + weakly yellowish,
C-, KC+ rose (light orange), P+ orange, UV-.
TLC/HPLC: cortical atranorin (minor) and usnic acid (minor); medullary
protocetraric acid (major), conprotocetraric acid (trace), virensic acid (trace),
subvirensic acid (trace), and convirensic (minor).
PARATYPES— Brazil, Sao Paulo State, Municipality of Sao Sebastiao, Centro de Biologia
Marinha da USP, Ponta do Baleeiro, southward rocky shore, 23°48°S, 45°37W, 15 m
alt., granitic rock, leg. M.P. Marcelli 2215 (S), 2222 (H), 2228 (SP), 2236 (B), 2246 (SP),
19-III-1988 (SP).
COMMENTS—Parmotrema graniticum is a saxicolous species morphologically
similar to PR. mordenii (Hale) Hale in the flat habit and common subpustular
soredia, but with a very different chemistry.
In addition to the chemistry, P graniticum is characterized by its +flat thallus,
the eciliate margins, irregular linear or subcapitate soralia appearing at the.
apices of short marginal lacinules, grouped rhizines and weakly sublageniform
conidia. Unlike many other Parmotrema species with sorediate margins, the
lower distal portions of P graniticum are almost always brown (rarely black),
usually with a reddish tone, even on sublacinulate and/or sorediate lobes.
Parmotrema spp. nov. (Brazil) ... 241
Figure 3. The fragment constituents of the type of P graniticum before the isotype making (a
single specimen). Bar = 1 cm.
242 ... Marcelli, Benatti & Elix
Although morphologically similar to P mordenii and some saxicolous
specimens of P. praesorediosum (Ntl.) Hale, P graniticum differs in having a
more coriaceous thallus, in the cortical and medullary chemistry, and in the
smaller, sub-bacilliform conidia, which have almost imperceptible dilations
when compared to those of species with pronounced, sublageniform conidia.
Whereas P. graniticum contains cortical usnic acid and atranorin and the
protocetraric acid chemosyndrome in the medulla, P. praesorediosum (which
may also be corticolous), contains cortical atranorin and protopraesorediosic
acid, praesorediosic acid and unknown fatty acids in the medulla. Furthermore,
P. praesorediosum has a more delicate thallus with ascending lobe margins and
develops chiefly farinose soredia and subcontinuous to labriform marginal
soralia.
Parmotrema pardi (Krog & Swinscow) Krog & Swinscow, so far known only
from East Africa (Krog & Swinscow 1981, 1983; Swinscow & Krog 1988), is the
only other saxicolous species with sorediate lobes and medullary protocetraric
acid, but has larger (0.8-2.0 cm wide), generally entire or subcrenate rather
than sublacinulate lobes, sparse rhizines, imperforate apothecia that became
perforate at maturity, filiform 10-14 um conidia, only atranorin present at the
upper cortex and fumarprotocetraric acid as well as protocetraric acid in the
medulla.
Parmotrema gardneri (C.W.Dodge) Sérus. (Dodge 1959, Krog & Swinscow
1981, Swinscow & Krog 1988) is a further coriaceous, marginally sorediate,
eciliate Parmotrema species which contains medullary protocetraric acid.
However, it is a corticolous species with wider lobes (8.0—20.0 mm vs. 1.5—7.0
mm wide), occasionally develops short cilia and has larger ascospores than
those of P. graniticum (16-26 x 8-14 um vs. 7.5-12.5 x 4-—6.5 Mm). Further, the
upper cortex of P. gardneri contains only atranorin and chloroatranorin. [NB.
medullary protocetraric, subvirensic, virensic, convirensic and conprotocetraric
acid are present in P. gardneri].
Further similar sorediate species containing the protocetraric chemo-
syndrome include, P dilatatum (Vain.) Hale, P schindleri Hale, P. fractum
(Hale) Hale, P ravum (Krog & Swinscow) Sérus., and P. dominicanum (Vain.)
Hale. However, these species exhibit significant differences as shown in
Table 1.
In the specimens studied, we observed that many young apothecia had
darkened discs without asci. The weak K+ yellowish medullary reaction
observed in parts of some specimens is believed to be due the presence of small’
amounts of atranorin in the medulla adjacent to the upper cortex.
The specific epithet refers to the substrate on which the species commonly
STOWS.
243
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Parmotrema milanezii Marcelli, Benatti & Elix, sp. nov. Fig. 4
MycoBAnk MB 511905
DiaGnosis: Species cum thallo simili Parmotrematis dilatati sed thallo membranaceo,
margine abundanter ciliato, ciliis plerumque ramosis etiam in soraliis formantibus
differt; atranorinam, chloroatranorinam, acidum usnicum, acidum protocetraricum,
acidum echinocarpicum, acidum conechinocarpicum, acidum convirensicum, acidum
subvirensicum et acidum secalonicum A continens.
Ho.otype-Brazil, Sao Paulo State, Municipality of Itanhaém, Jardim Grandesp, km 116
of Padre Manoel da Nobrega Highway, between the highway and the foot of the Serra
do Mar mountains, high restinga forest ca. 2 km from the highway, at the edge of track
following the direction of the Serra do Mar, 24°12’S, 46°53°W, 16 m alt., small tree
trunk, leg. M.P.Marcelli & J. Vieira Filho 9193, 30-01-1982 (SP; isotype in S).
THALLUS up to 20.5 cm diam., submembranaceous to membranaceous,
corticolous, greenish gray becoming pale brown in the herbarium, lobate; LoBEs
3.5-11.0(—20.0) mm wide, irregularly branched, younger lobes contiguous to
subimbricate, becoming crowded at maturity, adnate to weakly ascending when
sorediate, adnate to loosely adnate; apices +flat, rotund to subrotund; MARGIN
initially smooth becoming subcrenate, flat near the apices but becoming
subundulate in part and weakly ascending in older parts, becoming involute or
revolute when sorediate, entire to incised, +sublacinulate, ciliate; ci1a black,
simple to dichotomous or irregularly branched, 0.1—2.5(—3.8) x 0.05-0.1 mm,
usually abundant along the margins even amidst the soralia, absent at the lobes
apices.. UPPER SURFACE smooth to subrugose in part, distal lobes continuous,
becoming irregularly cracked within; maculae, pustules and isidia absent,
lacinules and soredia present; LACINULES short, irregular on the margins and
at the apices of older lobes, simple to rarely furcate, flat, 0.4-2.2 x 0.3-0.8 mm,
truncate, underside generally white, rarely variegated or black, less frequently
concolorous with the lower margin, rarely intermixed with small, irregular,
adventitious lobules; soRALIA marginal on small lacinules, irregularly linear
to subcapitate, often coalescing laterally and becoming +dense and irregular,
+raised, SOREDIA farinose to subgranular, coarse in part. MEDULLA white,
pigments absent. LowER suRFACE black, shiny, smooth to rugose, veined to
very rarely subpapillate, lacking rhizines in part, MARGIN shiny, brown, smooth
or rarely rugose, weakly veined to papillate, 1.5-6.5 mm wide, naked, becoming
cream coloured or variegated under sorediate lobes; RHIZINEs black, simple to
+furcate, 0.2-0.8(-—1.9) x 0.05-0.1 mm, absent to frequent in part, grouped.
APOTHECIA submarginal, very rare, concave, 2.3 mm diam., substipitate,
margin smooth, sorediate, eciliate, amphithecia and stipe smooth; pDisc
brown, epruinose, imperforate; AscosPoREs not found; pycNIDIA very rare,
submarginal, with black ostioles; CONIDIA not seen.
SPOT TESTS: upper cortex K+ yellow, UV-; medulla K+ yellow, C-, KC+ rose,
P+ orange, UV-.
Sa tel
Parmotrema spp. nov. (Brazil) ... 245
Figure 4. Type of P. milanezii before the making of the isotype (upper portion).
Bar = 1 cm.
246 ... Marcelli, Benatti & Elix
TLC/HPLC: cortical atranorin (minor), chloroatranorin (minor) and usnic
acid (trace); medullary protocetraric acid (major), echinocarpic acid (minor),
subvirensic acid (trace), convirensic acid (trace), conechinocarpic acid (+trace)
and secalonic A acid (+trace).
ParRATYPES— Brazil, Sao Paulo State, Municipality of Ilha Comprida, southern part of
the island, at the edge of the road near the raft to Cananéia, low restinga wood, 25°01’S,
47°54’W, 6 m alt., on tree branch inside the wood, leg. L.S.Canéz, A.A.Spielmann,
M.N.Benatti & M.P.Marcelli 1291, 02-IV-2004 (NY); idem, bush branch inside the wood,
leg. L.S.Canéz, A.A.Spielmann, M.N.Benatti & M.P.Marcelli 1312, 02-IV-2004 (SP).
COMMENTS- Parmotrema milanezii is distinguished by its flat, adnate,
membranaceous thallus, the lower cortex with sparse groups of rhizines, the
sorediate margins with small lacinules, and the dense, branched cilia within
the soralia, each cilium (usually) fixed to an individual soredium grain, which
can be readily detached by removing the cilium. This feature distinguished it
from all other sorediate Parmotrema species, where the cilia are usually not
present at the margin where it begins to open or erode to form soredia. ‘This
species contains the protocetraric and echinocarpic acid chemosyndromes in
the medulla, thus resembling the chemistry of P dilatatum.
Like P. milanezii, P dilatatum has sorediate margins with small lacinules
but lacks the very dense, commonly branched cilia arising from individual
grains of soredia. Further, P dilatatum is normally described as eciliate (e.g.
Vainio 1890, Swinscow & Krog 1988, Louwhoff & Elix 1999, Elix 1994), or
cited as producing very sparse simple, short marginal cilia, which are generally
restricted to the lobes axils (see Hale 1965, Fleig 1997).
Parmotrema perlatum (Huds.) M.Choisy, another K+ yellow ciliate and
sorediate species, usually has simple cilia, which do not occur in the sorediate
parts and it contains the stictic acid chemosyndrome in the medulla. Further, P.
perlatum has narrower lobes, shorter cilia (rarely more than 1.5 mm long), and
a more densely rhizinate lower surface with scattered rhizines.
The overall morphology of P. schindleri is similar to P milanezii, but it
differs in having short, sparse cilia in the axils, sparse marginal lacinules and
in containing protocetraric and gyrophoric acids in the medulla (Hale 1986).
P. schindlerii was recorded as a strictly saxicolous species.
This new species is named in honor of the Brazilian mycologist Dr. Adauto
Ivo Milanez, a distinguished mycologist and member of the Brazilian scientific
community who has expended considerable effort in encouraging Manisatohoyaice
research at the Instituto de Botanica.
Parmotrema spp. nov. (Brazil) ... 247
Acknowledgements
The authors wish to thank Harrie J.M. Sipman (Berlin) and Richard Harris (New York)
for the critical revision of the manuscript, and Teuvo Ahti (Helsinki) for help with the
Latin diagnosis.
This work could not have been accomplished without the support of the Conselho
Nacional de Desenvolvimento Cientifico e Tecnolégico (CNPq) for a masters scholarship
to the second author and for research support to the first author.
Literature cited
Benatti MN. 2005. Os géneros Canomaculina, Parmotrema e Rimelia (Parmeliaceae, Ascomycetes)
no litoral centro-sul do Estado de Sao Paulo. Mastership dissertation, Instituto de Botanica,
Sao Paulo, 389 p.
Brodo IM, Sharnoff SD, Sharnoff S. 2001. Lichens of North America. Yale University Press. New
Haven e London. 795 p.
Bungartz F. 2001. Analysis of lichen substances. In http://gila-asu.edu/ViewCVS/ viewcvs.cgi/
*checkout*/asulichens/plb400/laboratory/chemistry/tlc.html. Accessed on December 2007.
Dodge CW. 1959. Some lichens of tropical Africa. III. Parmeliaceae. Annals of the Missouri
Botanical Garden 46: 39-193.
| Donha CG. 2005. Os géneros Canomaculina, Parmotrema e Rimelia (Ascomycota Liquenizados)
na Area de Protecao Ambiental de Guaraquecaba, Parana, Brasil. Dissertacao de Mestrado.
Universidade Federal do Parana. 118 p.
Eliasaro S. 2001. Estudio taxonomico y floristico sobre las parmeliaceae sensu stricto (Ascomycota
liquenizados) del Segundo Planalto del Estado de Parana, Brasil. Tese de Doutorado. Facultad
de Ciencias Exactas y Naturales, Universidad de Buenos Aires.
Elix JA 1994. Parmotrema. In Flora of Australia, Lichens. Introduction, Lecanorales 2. vol. 55:
140-162. Australia Government Publishing Service, Canberra.
Elix JA, Giralt M, Wardlaw JH. 2003. New chloro-depsides from the lichen Dimelaena radiata.
| Bibliotheca Lichenologica 86: 1-7.
| Fleig M. 1997. Os géneros Parmotrema, Rimelia e Rimeliella (Lichenes - Ascomycota, Parmeliaceae)
no Rio Grande do Sul, Brasil. Tese de Doutorado, Instituto de Biociéncias da Universidade de
| Sao Paulo. 250 p.
| Hale ME. 1965. A Monograph of the Parmelia subgenus Amphigymnia. Contributions from the
United States National Herbarium 36(5): 193-358.
| Hale ME. 1986. New species in the lichen Family Parmeliaceae (Ascomycotina). Mycotaxon 25:
| 85-93.
Krog H, Swinscow TDV. 1981. Parmelia subgenus Amphigymnia (lichens) in East Africa. Bulletin
| of British Museum of Natural History (Bot.) 9(3): 143-231.
| Krog H, Swinscow TDV. 1983. A new species and new combinations of Parmotrema (Parmeliaceae).
Lichenologist 15(2): 127-130.
Louwhoff SHJJ, Elix JA. 1999. Parmotrema and allied lichen genera in Papua New Guinea.
| Bibliotheca Lichenologica 73: 1-152.
| Marcelli MP. 2004. Checklist of lichens and lichenicolous fungi of Brazil. Version 1: June 2004.
| http://www. biologie.uni-hamburg.de/checklists/brazil_l.htm.
Nash TH II, Elix JA. 2004. Parmotrema. In Nash III, TH, Ryan, BD, Diederich P, Gries C, Bungartz F
(Eds). Lichen flora of the greater Sonoran Desert Region. Volume 2. Lichens Unlimited, Arizona
State University, Tempe, Arizona, USA. pp. 318-329.
248 ... Marcelli, Benatti & Elix
Swinscow TDV, Krog H. 1988. Macrolichens of East Africa. British Museum of Natural History.
London. 390 p.
Vainio EA. 1890. Etude sur la classification naturelle et la morphologie des Lichens du Brésil, pars
prima. Acta Societatis pro Fauna et Flora Fennica 7(1): i-xxix, 1-247.
Vainio EA. 1896. Lichenes Antillarum a W.R. Elliot collecti. The Journal of Botany 34: 31-36.
MYCOTAXON
Volume 105, pp. 249-256 July-September 2008
New records of three freshwater ascomycetes
from an urban lagoon of Tabasco, Mexico
EDMUNDO ROSIQUE-GIL’, MARIA C. GONZALEZ! & JOAQUIN CIFUENTES?
*erosique@ibiologia.unam.mx
‘Departamento de Botanica, Instituto de Biologia, Universidad Nacional Auténoma de
México, Ciudad de México, DF 04510, México
*Herbario FCME, Facultad de Ciencias, Universidad Nacional Autonoma de México AP
| 70-399, Coyoacan México DF 04510, México
Abstract — A survey of lignicolous freshwater ascomycetes was carried out in Las
Ilusiones Lagoon located in the metropolitan area of Villahermosa City, State of Tabasco,
Mexico. Aniptodera inflatiascigera, Ascosacculus aquaticus and A. heteroguttulatus are
recorded for the first time from Mexico. This study is an important contribution to the
knowledge of the worldwide distribution of these three species from tropical freshwater
ecosystems.
Key words — Halosphaeriales, lentic habitat, Sordariales, submerged wood panels,
taxonomy
| Introduction
:
|
| The number of fungal species described worldwide is around 80,060 (Kirk et al.
2001). Of these, only about 500 species are freshwater meiosporic euascomycetes
| (Tsui & Hyde 2003, Vijaykrishna et al. 2006, Shearer et al. 2007). Little is known
jabout the biodiversity of this group of ascomycetes from tropical eco-regions
(Vijaykrishna & Hyde 2006) and particularly Mexico. The only knowledge of
| ascomycetes from a lentic freshwater environment is from a study by Gonzalez &
| Chavarria (2005) of freshwater ascomycetes from a temperate urban ecosystem
|situated in the metropolitan zone of Mexico City.
| During the past two years a biodiversity project to study lignicolous
ees ascomycetes of the State of Tabasco was undertaken because it is
|located in one of the highest precipitation regions of Mexico, which results in
jan elevated diversity of natural and urban lentic habitats. Las Ilusiones Lagoon
is located in the residential section of Villahermosa City, capital of the State of
| Tabasco. This lotic environment has vital importance as a water reservoir for the
city and it is a popular recreational area. In this study, we identify and illustrate
|three freshwater ascomycetes recorded from Mexico for the first time.
|
250 ... Rosique-Gill, Gonzalez & Cifuentes
Materials and methods
Wood panels submerged for three months in the Las Ilusiones Lagoon
(17°59°22”, 18°01722” N and 92°55720", 92°56'40” W, cover surface 198.4
ha), located in the urban zone of Villahermosa City, Tabasco State, México
(Rodriguez 2002). The phosphorous and nitrogen elevated values registered
at Las Ilusiones Lagoon suggest eutrophication, based on boundary values
of the trophic classification system (OECD 1982). The exposed panels were
retrieved, placed in Zip-lock plastic bags, transported to the laboratory in a
cooler containing ice and processed within 6 h. Collected material was washed
in tap water and placed in transparent plastic boxes containing moist paper
towels (Jones 1971). Samples were incubated at laboratory conditions (25-30
C) under natural daylight. Wood panels were examined at one month for the
presence of fungal fruiting structures. For morphological examination, squash
mounts of ascomata in sterile water were prepared on glass slides, and ascomata, ©
paraphyses, asci and ascospores were measured and photographed using an»
Olympus BX50 light/interference/phase contrast microscope. Dried specimens
on wood panels and permanent microscope slides, prepared according to the -
double cover glass method (Volkmann-Kohlmeyer & Kohlmeyer 1996), have -
been deposited in the fungal collection of Herbario Nacional (MEXU) of the ©
Institute of Biology, Universidad Nacional Autonoma de México. |
Taxonomy |
Aniptodera inflatiascigera K. M. Tsui, K.D. Hyde & Hodgkiss, |
Sydowia 49: 188 (1997). Figs. 1-6 —
MATERIAL EXAMINED — MEXICO. TasBasco: Centro Municipality, LAs ILUSIONEs
LAGOON (92°55'75” W 18°00’57”N) 24 Apr 2006, E Rosique-Gil, MC Gonzalez, J
Cifuentes MGFW5 and slide 5 (MEXU 25351, Pinus sp. wood test block submerged 3 mo
and incubated 1 mo.
Aniptodera inflatiascigera is very similar in ascospore form to other species,
such as A. fusiformis, A. limnetica and A. margarition, in having hyaline, bi-
celled thick-walled ascospores. However, A. inflatiascigera has larger ascospores
(32-46 x 15-22 um) compared to those of A. fusiformis (19-28 x 8-12 um), A.
limnetica (18-24 x 8-11 um), and A. margarition (15-22 x 9-13 um) (Shearer
1989, Tsui et al. 1997, Hyde et al. 1999). Also, A. inflatiascigera resembles A.
chesapeakensis and A. lignatilis in the conspicuous retraction of cytoplasm in
the ascus apices (Shearer & Miller 1977, Hyde 1992). The large asci that swell on
release from the ascomata is an exclusive characteristic of A. inflatiascigera. The —
features of the Mexican specimen, however, agree with those of A. inflatiascigera,
particularly the thick-walled ascospores (29-30 x 10-15 um), the swollen asci,
the thickened ascus apex and the subapical retraction of cytoplasm in the ascus.
New freshwater ascomycetes for Tabasco (Mexico) ... 251
Figs. 1-6. Aniptodera inflatiascigera. 1. Ascoma on wood. 2. Conical, periphysate, hyaline neck
with ascospores in canal. 3. Swollen asci. 4. Immature ascus and catenophyses. 5. Asci in various
stages of development showing the apical thickening and cytoplasm retraction below the apex.
6. Thick-walled ascospores.
Bars 1 = 100 pm, 2 = 20 um, 3, 4, 5 = 50 um, 6 = 20 um. Photomicrographs | taken with
light microscopy, 2, 3, 6 with interference contrast microscopy and 4, 5 with phase contrast
microscopy.
. Rosique-Gill, Gonzalez & Cifuentes
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New freshwater ascomycetes for Tabasco (Mexico) ... 253
Figs. 11-16. Ascosacculus heteroguttulatus. 11. Ascomata on wood. 12. Accumulation of ascospores
at apex of ascomata. 13. Subglobose pigmented ascoma with hyaline neck. 14. Catenophyses.
15. Asci in various stages of development. 16. Ascospores with distinctive two guttules in one cell
and numerous small guttules in the second cell.
Bars 11 = 500 um, 12 = 100 pm, 13 = 500 pm, 14 = 10 um, 15 = 20 um, 16 = 10 um.
Photomicrographs 11, 12, 13 taken with light microscopy, 14, 15 with phase contrast microscopy
and 16 with interference contrast microscopy.
254 ... Rosique-Gill, Gonzalez & Cifuentes
In this study, this species occurred rarely on the submerged panel wood at lentic
and heavily organic polluted urban environment. Aniptodera inflatiascigera
is an uncommion and new ascomycete for Mexico. Known distribution from
Brunei, Hong Kong and Philippines in lotic ecosistems (Tsui et al. 1997, Ho et
al. 2001).
Ascosacculus aquaticus (K.D. Hyde) J. Campb., J.L. Anderson & Shearer, Mycologia
95: 545 (2003). Figs. 7-10
MATERIAL EXAMINED — MEXICO. Tasasco: Centro Municipality, Las ILUSIONEs
LaGoon (92°55’51” W 18°00°39”N) 24 Apr 2006, E Rosique-Gil, MC Gonzalez, J
Cifuentes MGFWé6 and slide 6 (MEXU 25352, Tabubeia sp. wood test block submerged
3 mo and incubated 1 mo.
Ascosacculus aquaticus has subglobose or pyriform 180-205 x 140-164 um
ascomata, immersed or semi-immersed, light brown or brown, membranaceous,
ostiolate, with a neck, solitary or gregarious; long or short neck, up to 320
um long, 30-42 um diam, hyaline and periphysate; catenophyses present;
saccate, 56 x 30 um, 8-spored, thin-walled, with no apical thickening or pore,
deliquescing early asci; fusiform or cylindrical ascospores, 33.5-64 x 7-10 um, |
hyaline, bi-celled, sub-medial septa, thin-walled, highly guttulate, with hamate
appendages at each end (Hyde 1992). The Mexican material has fusiform or
cylindrical hyaline, bi-celled, sub-medial septa, thin-walled, highly guttulate,
with hamate appendages at each end, but smaller ascospores (22-32 x 7-8
um). However, the Mexican specimen is considered to be conspecific with
A. aquaticus. This is the first record of A. aquaticus from lentic and heavy
organic polluted freshwater environment from Mexico. Known distribution
only of lotic ecosystems from Australia (Hyde 1992) and Costa Rica (Campbell
et al. 2003).
Ascosacculus heteroguttulatus (S.W. Wong, K.D. Hyde & E.B.G. Jones)
J. Campb., J.L. Anderson & Shearer, Mycologia 95: 545 (2003). Figs. 11-16
MATERIAL EXAMINED — MEXICO. TasBasco: Centro Municipality, Las ILUSIONEs
LAGOON (92°56'28" W 18°00°85"N) 4 Dec 2006, E Rosique-Gil, MC Gonzalez, J
Cifuentes MGFW7 and slide 7 (MEXU 25353, Pinus sp. wood test block submerged 3 mo
and incubated 1 mo.
Ascosacculus heteroguttulatus has black, globose to subglobose, 112-168 um
diam ascomata, immersed or superficial, with long neck; catenophyses present;
early deliquescent 8-spored asci; ellipsoid 27-37 x 9-17.5 um ascospores, 2-
celled, hyaline, apical cell with one or two large lipid guttules, basal cell with
numerous small guttules (Wong et al. 1998, Hyde et al. 1999, Anderson &
Shearer 2002). The characteristics of the Mexican material agree with those of
the protologue, particularly, the guttulate pattern of the ascospores and their
size (20-27 x 7-9 um). This species is new for Mexico and the only known
New freshwater ascomycetes for Tabasco (Mexico) ... 255
distribution in Americas is from Florida, USA (Anderson & Shearer 2002)
and Costa Rica (Campbell et al. 2003). Although this species was originally
reported from Australia, Brunei, Hong Kong, Mauritius, Philippines and South
Africa, the subsequent Americas records suggest a possible worldwide tropical
and subtropical distribution of this fungus. Interestingly, the Mexican record
of A. heteroguttulatus is the first from a heavily organic polluted urban body
of water.
Acknowledgments
The authors are grateful to the reviewers of the manuscript Dr. Carol A. Shearer, Kevin
D. Hyde and Shaun Pennycook. Also, the authors thank Biol. Andrés Pedrero Sanchez,
Director de Recursos Naturales de la Secretaria de Desarrollo Social y Proteccién al
Ambiente del Estado de Tabasco for field exploration permission and for providing
a boat for transportation. This work was supported by grant of Consejo Nacional de
Ciencia y Tecnologia (CONACYT-SEP clave U1 60502) to Instituto de Biologia, UNAM.
The first author thanks the Consejo Nacional de Ciencia y Tecnologia for a fellowship
to obtain the Doctor of Sciences degree (CONACYT 163202) and the Posgrado en
Ciencias Bioldgicas, Universidad Nacional Aut6noma de México.
Literature cited
Anderson JL, Shearer CA. 2002. Halosarpheia heteroguttulata: anamorph and report from the
northern hemisphere. Mycotaxon 82: 115-120.
Campbell J, Anderson JL, Shearer CA. 2003. Systematics of Halosarpheia based on morphological
and molecular data. Mycologia 95: 530-552.
Gonzalez MC, Chavarria A. 2005. Some freshwater ascomycetes from Mexico. Mycotaxon 91:
315-322.
Ho WH, Hyde KD, Hogdkiss IJ, Yanna. 2001. Fungal communities on submerged wood from
streams in Brunei, Hong Kong, and Malaysia. Mycol Res 105: 1492-1501.
Hyde KD. 1992. Tropical Australian freshwater fungi. IV. Halosarpheia aquatica sp. nov.,
Garethjonesia lacunosispora gen. & sp. nov. and Ophioceras dolichostomum (Ascomycetes). Aust
Syst Bot 5: 407-414.
Hyde KD, Ho W-H, Tsui CKM. 1999. The genera Aniptodera, Halosarpheia, Nais and Phaeonectriella
from freshwater habitats. Mycoscience 40: 165-183.
Jones EBG. 1971. Aquatic fungi. In Booth C ed, Methods in microbiology. vol 4, Academic Press,
London pp 335-365.
Kirk PM, Cannon PF, David JC, Stalpers JA. 2001. Ainsworth & Bisby’s Dictionary of the fungi. 9th
ed. Cab International, Wallingford.
OECD [Organization for Economic Cooperation and Development]. 1982. Eutrophication of
waters, monitoring, assessment and control. OECD cooperative programme on monitoring of
inland waters (eutrophication control). Directorate OECD, Paris.
Rodriguez RE. 2002. Las lagunas continentales de Tabasco. Universidad Juarez Auténoma de
Tabasco, México.
Shearer CA. 1989. Aniptodera (Halosphaeriaceae) from wood in freshwater habitats. Mycologia 81:
139-146.
256 ... Rosique-Gill, Gonzalez & Cifuentes
Shearer CA, Miller M. 1977. Fungi of the Chesapeake Bay and its tributaries. V. Aniptodera
chesapeakensis gen. et sp. nov. Mycologia 69: 887-898.
Shearer CA, Descals E, Volkmann-Kohlmeyer B, Kohlmeyer J, Marvanova L, Padgett D, Porter D,
Raja HA, Schmit JP, Thorton HA, Voglymayr H. 2007. Fungal biodiversity in aquatic habitats.
Biod Cons 16: 49-67.
Tsui KM, Hyde KD. 2003. Introduction. Fun Div Res Ser 10: 1-9.
Tsui KM, Hyde KD, Hodgkiss IJ. 1997. A new species of Aniptodera (Ascomycetes) from Hong
Kong and the Philippines. Sydowia 49: 187-192.
Vijaykrishna D, Hyde KD. 2006. Inter and intra stream variation of lignicolous freshwater fungi in
tropical Australia. Fungal Divers. 21: 203-224.
Vijaykrishna D, Jeewon R, Hyde, KD. 2006. Molecular taxonomy, origins and evolution of
freshwater ascomycetes. Fungal Divers. 23: 367-406.
Volkmann-Kohlmeyer B, Kohlmeyer J. 1996. How to prepare truly permanent microscope slides.
Mycologist 10: 107-108.
Wong SW, Hyde KD, Jones EBG. 1998. Halosarpheia heteroguttulata sp. nov. from submerged
wood in streams. Can J Bot 76: 1857-1862.
MY COTAXON
Volume 105, pp. 257-267 July-September 2008
New records and distribution of rust fungi from Pakistan
N. S. AFSHAN, A. N. KHALID & A. R. NIAZI
pakrust@gmail.com
Department of Botany, University of the Punjab
Quaid-e-Azam Campus, Lahore, 54590, Pakistan
Abstract — A contribution is made to the rust fungus flora of Pakistan with Uromyces
dactylidis, U. graminicola and U. peglerae var. peglerae reported as new records for the
country. New observations on previously known species are also noted. The spermogonia
(0) of Aecidium clematidis and A. montanum, spermogonia (0) and aecia (I) of Puccinia
prenanthis var. himalensis are described for the first time. Phragmidium rosae-moschatae
on Rosa webbiana var. microphylla, Puccinia levis var. panici-sanguinalis on Digitaria
radicosa, P. conclusa on Cyperus iria, and Uromyces setariae-italicae on Setaria viridis
are new hosts for rust fungi in Pakistan.
Key words — biodiversity, Melampsora epitea, Uredinales, vegetation
Introduction
This paper is a continuation of the enumeration of the rust fungi from Pakistan
(Afshan et al. 2007, Afshan et al. 2008a, Afshan et al. 2008b, Afshan et al.
2008c). Until now, only 21 genera and approximately 400 species of rust fungi
are known from Pakistan (Ahmad et al. 1997, Afshan et al. 2007, Afshan et al.
2008). These figures indicate that the number of rust species collected from
this area is far less than expected in relation to the vegetation. Comprehensive
surveys of unexplored areas are needed to understand the biodiversity of
Uredinales of Pakistan.
In the present investigation, Uromyces dactylidis, U. graminicola and
U. peglerae var. peglerae are re-described and illustrated. These rust fungi
are new to our rust flora. The spermogonia (0) of Aecidium clematidis and
A. montanum, spermogonia (0) and aecia (I) of Puccinia prenanthis var.
himalensis are reported for the first time from Pakistan. New hosts for Pakistan
include Clematis montana for Aecidium clematidis, Cyperus iria for P. conclusa,
Digitaria radicosa for P. levis var. panici-sanguinalis, Panicum antidotale for
Uromyces graminicola, Rosa webbiana var. microphylla for Phragmidium rosae-
moschatae and Setaria viridis for Uromyces setariae-italicae. New observations
258 ... Afshan, Khalid & Niazi
on previously known species were made. Melampsora epitea, Phragmidium
shogranense, Puccinia apludae, P. cirsii, P. helianthi, P. sorghi, Uromyces clignyi,
U. ignobilis and U. orientalis, already known from Pakistan, are reported from
new localities.
Materials and methods
Free hand sections of infected portions of material and spores were mounted
in lactophenol and gently heated to boiling. The preparations were observed
under a NIKON YS 100 microscope. Drawings of spores and paraphyses were
made using a camera lucida (Ernst Leitz, Wetzlar, Germany). Spore dimensions
were measured using an ocular micrometer. At least 25 spores were measured
for each spore stage.
Enumeration of taxa
1. Uromyces dactylidis G. H. Otth, Mitt. naturf. Ges. Bern: 85 (1861). (Fig. 1)
Spermogonia and aecia not seen. Uredinia on abaxial leaf surface brown,
minute. Urediniospores globose to ovoid or broadly ellipsoid, 20-23 x 20-24
tum; wall 1.5-2 um thick, hyaline to pale brown, echinulate; germ pores 5-7(-9),
scattered, obscure; pedicel short, hyaline. Telia abaxial, mostly amphigenous,
dark brown to blackish brown, subepidermal, loculate with brownish
paraphyses. Teliospores ellipsoid to broadly ellipsoid or obovoid, (14—)16-23
x 20-26(-32) um; wall 1.5-2 um thick at side, 3-5 um thick apically, brown to
chestnut brown, smooth; apex mostly rounded, sometimes truncate; pedicel
hyaline to yellowish, collapsing, 5-8 x 40 um. |
MATERIAL EXAMINED: On Agrostis gigantea Roth, Pakistan, North Western Frontier
Province (NWFP), Khanspur, II + III, 24" May, 2006. # NSA 1706 (LAH).
Uromyces dactylidis is newly reported from Pakistan.
8s Saas
Fig. 1. Uromyces dactylidis
(A). Urediniospores showing germ pores and echinulate wall ornamentation (B). Teliospores.
Scale bar: 10 um.
2. Uromyces graminicola Burrill, Bot. Gaz. 9: 188 (1884). (Fig. 2)
Spermogoria and aecia not seen. Uredinia amphigenous or mostly on
adaxial surfac2, golden brown, minute. Urediniospores ellipsoid or obovoid
but mostly broadly ellipsoid, (19-)20-24 x (20-)24(-28) um; wall 1.5-2.5 um
Rust fungi new to Pakistan ... 259
D Te
Fig. 2. Uromyces graminicola (C). Urediniospores (D). Teliospores. Scale bar: 10 um.
thick, pale brown to golden brown, echinulate; germ pores 3-4, equatorial.
Telia amphigenous, dark brown to blackish brown, early exposed. Teliospores
ellipsoid to broadly ellipsoid or ovoid to obovoid, sometimes angular, 13-21 x
(20-)22-30 um; wall 1.5-2 um thick at sides, 5-10 um thick at apex, golden
brown to chestnut brown, smooth; apex mostly rounded, sometimes conical,
chestnut brown; pedicel hyaline to golden, persistent, 5-8 x 70-95 um.
MATERIAL EXAMINED: On Panicum antidotale Retz., Pakistan, North Western Frontier
Province (NWFP), Khanspur, II + III, 20" August, 2006. # NSA 2008 (LAH).
Uromyces graminicola is newly reported from Pakistan.
3. Uromyces peglerae Pole-Evans ex Syd. & P. Syd. var. peglerae,
Ann. Mycol. 12: 263 (1914). (Fig. 3)
Spermogonia, aecia and telia absent. Uredinia abaxial, mostly on adaxial leaf
surface, brown, 0.4-0.6 x 0.3-0.5 mm. Urediniospores globose to broadly
ellipsoid or ovoid, 18-22 x 21-27 um; wall thickness 1.5-2.0 um, yellowish
brown to dark brown, echinulate; germ pores 6-8, scattered or tending to be
bizonate; pedicel hyaline, 6-8 x 45-50 um.
Fig. 3 (E). Urediniospores of Uromyces peglerae var. peglerae. Scale bar: 10 um.
260 ... Afshan, Khalid & Niazi
MATERIAL EXAMINED: On Digitaria adscendens (Kunth) Henrard (= D. ciliaris (Retz.)
Koeler), Pakistan, Punjab, Lahore, II, 28" February, 2006. # NSA 2802 (LAH).
Puccinia digitariae-velutinae Vienn.-Bourg. has been reported on Digitaria
ciliaris from Karachi (Ahmad et al. 1997). Cummins (1971) reported Uromyces
peglerae var. peglerae on Digitaria spp. This rust is a new record for Pakistan.
4, Aecidium clematidis DC., in Lamarck & de Candolle,
Fl. frang., Edn 3 (Paris) 2: 243 (1805). (Fig. 4)
Spermogonia adaxial, close to aecia, brown, in small groups, subepidermal,
sunken in tissues of leaf, brown, 0.2-0.3 x 0.3-0.5 mm, with projecting ostiolar
paraphyses, minute. Aecia abaxial, orange, in groups, cup-shaped, 0.1-0.2 x
0.2-0.3 mm. Aeciospores hyaline, ellipsoid to globose, 17-22(-24) x 17-26
um, verrucose. Peridial cells hyaline, rhomboidal, 13-17(-24) x 16-24(-31)
um, verrucose.
MATERIAL EXAMINED: On Clematis montana Buch.-Ham., Pakistan, North Western
Frontier Province (NWFP), Ayubia, 0 + I, 24" May, 2006. # NSA 2406 (LAH).
Ahmad (1956a, b) reported A. clematidis on Clematis spp. from Gilgit. The
spermogonial stage of A. clematidis is herein described for the first time from
Pakistan.
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Fig. 4. Aecidium clematidis (F). Aeciospores (G). Peridial cells. Scale bar: 10 um.
5. Aecidium montanum E. J. Butler, Indian Forester: 30 (1905). (Fig. 5)
Spermogonia adaxial, close to aecia, in small groups, subepidermal, sunken
in leaf tissues, brown, 0.2-0.5 x 0.3-0.5 mm. Aecia abaxial, on yellowish orange
spots with brown margins, in groups or scattered, localized, 0.1-0.4 x 0.2-0.41
mm, cylindrical or cupulate. Aeciospores hyaline to yellow, 15-21 x 24-28 um,
verrucose. Peridial cells hyaline, rhomboidal, 15-23 x 23-27 um, verrucose.
MATERIAL EXAMINED: On Berberis lycium Royle, Pakistan, North Western Frontier
Province (NWFP), Mukshupuri track, 0 & I, 23"! May, 2006. # NSA 2306 (LAH).
Rust fungi new to Pakistan ... 261
Een
\ Moe,
Fig. 5. Aecidium montanum (H). Aeciospores (1). Peridial cell. Scale bar: 10 um.
Aecidium montanum has been reported on Berberis pachyacantha Koehne,
B. zabeliana C. K. Schneid. and B. lycium from Chitral, Nathia galli, Changla
galli, Kaghan valley, Quetta and Danyor (Ahmad 1956a). The spermogonial
stage of this anamorphic rust is newly recorded for Pakistan.
6. Puccinia prenanthis var. himalensis Barclay,
Sci. Mem. Med. Off. Army India 6: 65 (1891). (Fig. 6)
Uredinia and telia absent. Spermogonia adaxial, scattered among aecia, white.
Aecia abaxial, orange, in groups, cupulate. Aeciospores hyaline, obovoid
to ellipsoid, 19-23(-25) x 21-25 um; wall 2.5-3.5 um thick, verrucose.
Peridial cells hyaline, irregular in shape, (10—) 12-18(-24) x 15-18(-23) um,
verrucose.
MATERIAL EXAMINED: On Lactuca brunoniana (Wall. ex DC.) C. B. Clarke, Pakistan,
North Western Frontier Province (NWFP), Ayubia, Mukshupuri track, 0 + I, 24'" May,
2006. # NSA 24061 (LAH).
The aecidial and spermogonial stages of P prenanthis var. himalensis on
L. brunoniana are herein reported for the first time from Pakistan. Ahmad
(1956a, b) reported the uredinial and telial stages of P prenanthis var. himalensis
on the same host from Kaghan valley, Shogran, Poonch and Changla galli.
sot
og ai ae tSaae
Fig. 6. Puccinia prenanthis var. himalensis (J). Peridial cells (K). Aeciospores. Scale bar: 10 um.
262 ... Afshan, Khalid & Niazi
7. Phragmidium rosae-moschatae Dietel, Hedwigia 44: 132 (1905). (Fig. 7)
Spermogonia, aecia and uredinia absent. Telia abaxial, black, 0.3-0.4 x 0.2-
0.4 mm. Teliospores dark brown, mostly 6-7 celled, but 8-celled is also present;
29-34 x 55-106 um, papilla 12-20 um high, wall thickness 3-4 um; pedicel u
to 125 um long. 7
MATERIAL EXAMINED: On Rosa webbiana g =
var. microphylla Crép., Pakistan, North / /}
Western Frontier Province (NWFP), Ld
Khanspur, II, 05" August, 2006, NSA # foo
50806 (LAH). i |
H
Rosa webbiana var. microphylla is
a new host for Phragmidium rosae-
moschatae from Pakistan. Earlier, Ph. |
rosae-moschatae was reported on Rosa |
brunonii Lindl., (= R. moschata Herrm.), =
R. lacerans Boiss. & Buhse, R. webbiana
Wall. ex Royle and R. centifolia L. from
Quetta, Murree, Changla Gali, Peshawar,
Tarnab (NWFP), Matiltan (Swat) and
Battakundi (Kaghan Valley) (Ahmad et
al. 1997). yi |
Fig. 7. (L). Teliospores of
Phragmidium rosae-moschatae. L
Scale bar: 10 um.
8. Puccinia levis var. panici-sanguinalis (Rangel) Ramachar & Cummins,
Mycopath. Mycol. appl. 25(1-2): 44 (1965). (Fig. 8)
Spermogonia and aecia unknown. Uredinia hypophyllous, scattered, on
leaf sheaths, in long linear rows, dark brown, 0.09-0.1 x 0.08-0.1 mm.
Urediniospores globoid , obovoid or broadly ellipsoid, 20-25 x 21-29 um,
echinulate, pale brown to cinnamon brown, wall 1.5-2 um thick at sides , 2-3
[um at the apex; germ pores 3-4, equatorial; pedicel hyaline and minute. Telia
amphigenous, blackish brown, 0.09-0.2 x 0.1-0.3 mm. Teliospores ellipsoid
to broadly ellipsoid, usually tending to be diorchidioid and often strongly so,
21-30 x 20-31 um, wall 2-3 um at sides while 4-7 um thick over the pores;
reddish brown to dark chestnut brown; germ pores obscure, scattered; pedicel
colorless, persistent, 4-7 xm wide and up to 170 um long
MATERIAL EXAMINED: On Digitaria radicosa (C. Presl) Miq., Pakistan, Punjab, Lahore,
SHI Mycological Herbarium # NSA 16. 23™ October, 2006 (LAH).
Rust fungi new to Pakistan ... 263
4
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Fig. 8. Puccinia levis var. panici-sanguinalis (M).
Urediniospores (N). Ellipsoid to diorchidioid type of teliospores.
Scale bar: 10 um.
Ahmad (1956a, b) reported Puccinia levis var. panici-sanguinalis as Diorchidium
digitariae S. Ahmad on Digitaria bicornis (Lam.) Roem. & Schult. from Lahore.
No rust fungus had previously been reported on D. radicosa from Pakistan. So
D. radicosa is a new host for this rust species in Pakistan.
9. Puccinia conclusa Thim.,
Journal Sci. math. phys. nat. Lisboa, 1 Ser. 6(no. 24): 237 (1878). (Fig. 9)
=Puccinia romagnoliana Maire & Sacc., Ann. Mycol. 1: 220 (1903).
Spermogonia and aecia unknown. Uredinia hypophyllous, brown, 0.2-0.3 x
0.09-0.1 mm. Urediniospores globose to subglobose or obovoid, brown, 18-21
x 21-28 um, echinulate, wall 1.5-3 um thick; germ pores 2, equatorial; pedicel
hyaline, short. Telia hypophyllous, covered by epidermis, black, 0.2 x 0.1 mm,
with brown paraphyses. Teliospores ellipsoid to clavate or oblong, pale brown,
slightly constricted at septa, base attenuated, 13-21 x 44-60 um; apex rounded
or conical, 4—8 um thick; wall 1-2 um thick, smooth; pedicel pale brown, short,
up to 40 um long.
MATERIAL EXAMINED: On Cyperus iria L., Pakistan, Punjab, Lahore. SHI Mycological
Herbarium, # NSA 17. 25" October, 2006 (LAH).
Puccinia conclusa has been reported under the name P romagnoliana on
Cyperus difformis L. and C. rotundus L. from Chuharkana (Sheikhupura) and
Karachi by Ahmad (1956a, b), Ghaffar & Kafi (1968) and Hasnain et al. (1959)
as listed in Ahmad et al. (1997). Ahmad (1956a, b) reported P. philippinensis
Syd. & P. Syd. and P. cypericola S. Ahmad on Cyperus rotundus. No rust species
has previously been reported on Cyperus iria from Pakistan. So C. iria is also a
new host for P. conclusa in Pakistan.
264 ... Afshan, Khalid & Niazi
Lp omni
Fig. 9. Puccinia conclusa
(O). Urediniospores and paraphyses (P). Ellipsoid to clavate teliospores. Scale bar: 10 um.
10.Uromyces setariae-italicae Yoshina, Bot. Mag. Tokyo. 20: 247 (1906). (Fig. 10)
Spermogonia and aecia not seen. Uredinia amphigenous, scattered or in
rows, pulverulent, brown, 0.09-0.1 x 0.1-0.3 mm. Urediniospores subglobose
to ovoid or ellipsoid, brown to yellowish brown, echinulate, 23-27.5 x 23-29
um; wall 1.5-2.5 um thick; germ pores up to 4, equatorial; pedicels hyaline,
up to 40 um long. Telia amphigenous, covered by the epidermis, blackish,
small and inconspicuous, striated, 0.05-0.2 x 0.09-0.2 mm. Teliospores one
celled, variable, mostly angularly globoid or obovoid, 16-24 x 19-28 um, wall |
uniformly 1-2 um thick, smooth, yellowish brown to chestnut brown; apex not
thickened, rounded or truncate. Pedicel hyaline, collapsing, 4 x 15 um.
MATERIAL EXAMINED: On Setaria viridis (L.) P. Beauv., Pakistan, North Western Frontier
Province (NWFP), Khanspur, II + HI, 15"*September, 2006. # NSA 1509 (LAH).
|
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Fig. 10. Uromyces setariae-italicae
(Q) Urediniospores with a paraphysis. (R) Teliospores. Scale bar: 10 um.
“t
Rust fungi new to Pakistan ... 265
Ahmad (1956a, b) reported Uromyces setariae-italicae on Urochloa reptans (L.)
Stapf as Brachiaria reptans (L.) C.A. Gardner & C.E. Hubb. and S. glauca from
Lahore, Sheikhupura and Tandojam. Setaria viridis is a new host for Uromyces
setariae-italicae in Pakistan.
11. Melampsora epitea Thiim., Mitth. forst. Versuchs. Oster. 2: 38 (1879).
Melampsora epitea is reported for the first time from Dunga galli. It is already
reported on leaves of Salix spp., and S. tetrasperma from Kaghan valley, Batal,
Swat and Hasanabad by Ahmad (1956a, b), Malik & Virk (1968) and Ahmad &
Arshad (1972); from Choa Saiden Shah by Khalid et al. (1993a); on S. flabellaris
Andersson, from Kaghan valley by Ono (1992).
MATERIAL EXAMINED: On Salix tetrasperma Roxb., Pakistan, North West Frontier
Province (NWFP), Bhoorban, Dunga Galli, II. Aug. 10, 2006. NSA # 1008 (LAH).
12. Phragmidium shogranense Petr., Sydowia, 8: 162 (1954).
This rust has been previously reported on Rubus sp. from Kaghan valley and
Swat by Ahmad (1956a, b), additional hosts include R. pedunculosus D. Don
and R. pungens Cambess. (Khalid et al. 1993b) and R. aff. biflorus Buch.-Ham.
ex Sm. from Kaghan valley (Ono 1992).
MATERIAL EXAMINED: On Rubus sp., Pakistan, North West Frontier Province (NWFP),
Khanspur, II + II, 10™ Sep., 2006. NSA # 1009 (LAH).
13. Puccinia apludae Syd. & P. Syd. Annls mycol. 4: 436 (1906).
Puccinia apludae was previously reported on leaves of Apluda mutica L.
from Charsadda by Ahmad & Arshad (1972) but is herein reported for first
time from Khanspur (NWFP).
MATERIAL EXAMINED: On Apluda aristata L., Pakistan, North West Frontier Province
(NWFP), Khanspur, II+III, June 23, 2006. NSA # 2305 (LAH).
14. Puccinia cirsii Lasch, in Rabenhorst, Fungi europ. exsicc.: no. 89 (1859).
Ahmad (1956a, b) reported P. cirsii on Cnicus argyracanthus from Swat state
and Madian, on C. wallichii Hook. f. from Swat state, Kalam and Changla galli,
and on Cnicus sp. from Kaghan Valley and Shogran. It is an addition to the rust
fungi of Khanspur (NWEP).
MATERIAL EXAMINED: On Cnicus argyracanthus C. B. Clarke, Pakistan, North West
Frontier Province (NWFP), Khanspur, II, Aug. 10, 2006, NSA # 100806 (LAH).
15. Puccinia helianthi Schwein., Schr. naturf. Ges. Leipzig 1: 73 (1822).
Puccinia helianthi is reported for the first time from Lahore. It has been
previously reported on Helianthus annuus from Karachi, Peshawar, Rawalpindi,
Quetta and Tandojam (Ahmad et al. 1997).
266 ... Afshan, Khalid & Niazi
MATERIAL EXAMINED: On Helianthus annuus L., Pakistan, Punjab, Lahore, II, Aug. 9,
2006, NSA # 90806 (LAH).
16. Puccinia sorghi Schwein., Trans. Am. phil. Soc., n.s. 4(2): 295 (1832).
=Puccinia maydis Berenger, Atti Riunione Sci. Ital. 6 (Milano 1844): 475 (1845).
Puccinia sorghi has been previously reported on leaves of Zea mays from Quetta,
Kaghan, Kulali, Swat, Lahore, Karachi, Tandojam, Balakot, Naran, Sharan and
Shogran (Ahmad et al. 1997), but this is a new record for Khanspur, NWFP.
MATERIAL EXAMINED: On Zea mays L., Pakistan, North West Frontier Province (NWFP),
Khanspur, III, Aug. 23, 2006. NSA # 2307. (LAH).
17. Uromyces clignyi Pat. & Har., J. Bot. Morot 14: 237 (1900).
=Uromyces andropogonis-annulati Syd., P. Syd. & E.J. Butler, Annls mycol. 5(6): 492
(1907).
Ahmad (1956a) reported Uromyces clignyi from Gillanwali, Sangla Hill,
Rawalpindi, Kulali, Swat, Kalam, Karachi, and Gujranwala, but it is a new
record for Lahore.
MATERIAL EXAMINED: On Dichanthium annulatum (Forssk.) Stapf, Pakistan, Punjab,
Lahore, II, Sep. 20, 2006, NSA # 2009 (LAH).
18. Uromyces ignobilis (Syd. & P. Syd.) Arthur, Mycologia 7(4): 181 (1915).
This species has been previously reported on leaves of Sporobolus arabicus
from Lahore by Ahmad (1956a, b) and Masood et al. (1995), but this is the first
report on S. arabicus from Khanspur (NWEP).
MATERIAL EXAMINED: On Sporobolus arabicus Boiss., Pakistan, North West Frontier
Province (NWFP), Khanspur, II, May 23, 2006, NSA # 2305 (LAH).
19. Uromyces orientalis Syd. & P. Syd., Annls mycol. 5(6): 490 (1907).
Uromyces orientalis is reported for the first time from Lahore. It has been
previously reported on Indigofera linifolia from Sheikhupura, Zafarwal and
Gujrat by Ahmad (1956a, b, 1964a).
MATERIAL EXAMINED: On Indigofera linifolia (L. f.) Retz., Pakistan, Punjab, Lahore,
II+IH, Sep. 20, 2006. NSA # 20091 (LAH).
Rust fungi new to Pakistan ... 267
Acknowledgements
We sincerely thank Dr. Amy Rossman, Systematic Mycology and Microbiology
Laboratory, USDA-ARS, Beltsville and Dr. Mehrdad Abbasi, Iranian Research Institute
of Plant Protection, Department of Botany, Tehran, Iran, for their valuable suggestions
to improve the manuscript and acting as presubmission reviewers.
References
Afshan NS, Khalid AN, Niazi AR. 2008a. New records of graminicolous rust fungi from Pakistan.
Pak. J. Bot. 40(3): 1279-1283.
Afshan NS, Khalid AN, Javed H. 2008b. Further additions to the Rust flora of Pakistan. Pak. J. Bot.
40(3): 1285-1289.
Afshan NS, Berndt R, Khalid AN, Niazi AR. 2008c. New graminicolous rust fungi from Pakistan.
Mycotaxon 104: 123-130.
Afshan NS, Khalid AN, Abbasi M, Niazi AR. 2007. New records of rust fungi for Pakistan.
Mycotaxon 101: 233-237.
Ahmad S, Iqbal SH, Khalid AN. 1997. Fungi of Pakistan. Sultan Ahmad Mycological Society of
Pakistan, Department of Botany, University of the Punjab, Lahore, Pakistan.
Ahmad S. 1956a. Uredinales of West Pakistan. Biologia 2(1): 29-101.
Ahmad S. 1956b. Fungi of Pakistan. Biological Society of Pakistan, Lahore Monograph 1: 1-126.
Ahmad S. 1964a. Contributions to the fungi of Pakistan. IV. Biologia 10: 1-62.
Ahmad S, Arshad M. 1972. Contributions to the fungi of Pakistan. XII. Biologia 8: 7-17.
Cummins GB. 1971. The Rust Fungi of Cereals, Grasses and Bamboos. Springer Verlag, Berlin.
Heidelberg. New York.
Dade HA, Gunnell J. 1969. Class work with fungi. Commonwealth Mycological Institute Kew,
Surrey, England.
Ghaffar A, Kafi A. 1968. Fungi of Karachi. Pak. J. Sc. 20: 5-10.
Hasnain SZ, Khan A, Zaidi AJ. 1959. Rusts and smuts of Karachi. Bot. Deptt. Karachi Univ. Mont.
22 DD.30.
Hennen JF and McCain JW. 1993. New species and records of Uredinales from the Neotropics.
Mycologia 85: 970-986.
Khalid AN, Iqbal SH, Parveen B. 1993b. Rust flora of Pakistan. I. Genus Phragmidium Link., on
Rubus spp. Pakphyton 5: 133-136.
Malik SA, Virk. 1968. Contribution to the knowledge of parasitic fungi of Quetta-Kalat Region.
Biologia 14: 27-35.
Masood A, Khalid AN, Iqbal SH. 1995. New records of Graminicolous rust fungi (Uredinales) from
Pakistan. Sci. Int. (Lahore) 7(3): 415-416.
Ono Y. 1992. Uredinales collected in the Kaghan Valley, Pakistan. Cryptogamic flora of Pakistan
1; 217-240.
Sultan MA. 2005. Taxonomic study of rust flora of Northern Areas of Pakistan. Ph.D. Thesis,
University of the Punjab. Lahore, Pakistan.
MYCOTAXON
Volume 105, pp. 269-293 July-September 2008
Additions to the corticioids of the Caucasus
from NW Iran
MASOOMEH GHOBAD-NEJHAD!, NILS HALLENBERG?
& HEIKKI KOTIRANTA?
ghobadne@mappi.helsinki.fi, nils.hallenberg@dpes.gu.se heikki.kotiranta@ymparisto.fi
'Finnish Museum of Natural History, Botanical Museum
P.O.Box 7, FI-00014 University of Helsinki, Finland
*Dept of Plant and Environmental Institute, Goteborg University
Box 461, S-405 30 Goteborg, Sweden
°Finnish Environment Institute, Research Department
PO. Box 140, FI-00251 Helsinki, Finland
Abstract — A recent survey of wood-inhabiting Basidiomycota in NW Iran added 11
corticioids new to Iran, 6 of which are reported for the first time from the Caucasus
area. Additional finds of 10 rarely collected species in Iran are presented and data
on their occurrence in the Caucasus is also summarised. Most of the species are
illustrated and ecological and distribution notes are provided. Some rare or otherwise
noteworthy records are Hyphodontia erastii, Melzericium bourdotii, Peniophora laeta,
and Tubulicrinis incrassatus.
Key words — Arasbaran, Corticiaceae s.l.
Introduction
The Caucasus region, covering an area of some 532,700 km” between Black Sea
and Caspian Sea, comprises the countries Georgia, Armenia, Azerbaijan, parts
of European Russia, NE Turkey, and NW-N Iran. The montane forests of the
Arasbaran area and Talish forests in NW Iran, as well as the Hyrcanian forests
constitute the southern stretches of the Caucasus.
Intensive survey on wood-inhabiting Basidiomycota (polypores and
corticioids) in N Iran was carried out by Hallenberg during 1978-81 (Hallenberg
1978, 1979, 1980, 1981). Recent collections were made by Ghobad-Nejhad during
2005-06 in NW Iran, mainly in Arasbaran area. Situated in the northern part
of East Azerbaijan province, Arasbaran (Qara daq) covers an area of 900,000
hectares and belongs to the sub-humid zone in the northwest of Iran. The
vegetation is dominated by Quercus spp. and Carpinus betulus, accompanied
270 ... Ghobad-Nejhad, Hallenberg & Kotiranta
Azerbaijan
Armenia
Azerbaijan
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38°38" |
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: Province border
@@@ Arasbaran area
Fig. 1. Localities visited in Arasbaran area, NW Iran. For description of numbers see Table.
with Juniperus spp., Taxus baccata, Fraxinus rotundifolia, Lonicera spp., Acer
campestre etc. A large part of the Arasbaran area is covered by high peaks and
steep slopes. The area is affected from the west by mediterranean climate and
coincides in the south with Irano-Turanian steppes. Talish mountains in the
east connect the area to Hyrcanian forests, which have a markedly different
vegetation composition due mainly to their different amount of annual rainfall.
Arasbaran lacks some relict Hyrcanian elements such as Parrotia persica,
Quercus castaneifolia, etc. The area is threatened by over-grazing and logging
(e.g. Jalili et al. 2003).
Mycological inventories in the Arasbaran area are scanty, and none (if any) is
focused on wood-inhabiting Basidiomycota. These fungi, traditionally referred
to as corticioids and polypores, are distributed mainly in the orders Corticiales,
Gloeophyllales, Hymenochaetales, Russulales, Thelephorales, and Trechisporales,
all belonging to Agaricomycetes Doweld (sensu Hibbett et al. 2007).
This paper concentrates on new or some less collected corticioids from
NW Iran. Corticioids have for a long time been treated under the collective
family name Corticiaceae s.]. Larsson’s (2007) proposed set of fifty families to
accommodate the corticioid pee is still regarded as provisional and awaits
further modification.
New corticioids from the Iranian Caucasus ... 27 l
Table 1. Localities visited in NW Iran, E Azerbaijan province, and some of their
attributes. Numbers in the parentheses are shown in Fig. 1.
Locality Important vegetations Alt. (m) Coordinates
Jolfa, Missan (1) Acer monspessulanum, Paliurus spina- 820-1372 N38°5 1’: E46°26
christi, Carpinus betulus, Quercus sp.,
Juniperus excelsa
Jolfa, Daran (2) Juglans regia, Prunus armeniaca 1300 N38°39’; E4621
Shah-heidar (3) Juniperus sp., Paliurus spina-christi 700 N39°01?: E46°50°
Vaighan (4) Quercus sp., Carpinus betulus, Taxus 1350 N38°52’: E46°49”
baccata, Juniperus communis
Makidi valley (5) Quercus macranthera, Carpinus betulus, 1400 N38°50°: E46°55’
Acer campestre, Fraxinus exelsior,
Fraxinus rotundifolia
Kolashloo (6 cS 1820-18 sey RAeeeT
2 (6) Quercus sp., Carpinus betulus 2 .
Balan (7) Quercus sp., Acer sp., Lonicera spp., 2100 N 38°59”: E 47°29”
Viburnum lantana
Qale-darasi (8) — Quercus sp., Carpinus betulus 1800 N 38°50’: E47°00" ©
Kalale (9) Cornus mas, Carpinus betulus, Taxus 1200 N 38°56: E 46°45’
baccata, Acer sp., Quercus sp.
Oshtobin (10) _ Paliurus spina-christi, Amygdalus sp.. 1150 N38°50’: E46°29°
Quercus sp., Lonicera spp., Cornus mas,
____Cotinus coggygria -
Aynaloo (11) Quercus sp., Carpinus betulus, Acer 1750 N38°50°: E46°47’
campestre, Lonicera caucasica
Materials and methods
Material for this study was collected during two visits to Arasbaran area, in October-
November 2005 and 2006. Additional material collected from Russian Caucasus and NE
Turkey during 1989-96 is also added to the examined specimens in the text. Species are
treated alphabetically in two sections: first comes a list of new records, followed by the
list of some little collected corticioids. An asterisk (*) symbolises the new records to the
Caucasus, and a filled circle (°) marks records new to Iran. Old and recent accounts of
all species in the Caucasus, if applicable, are outlined and brief notes on substrates and
distribution are given. Specimens were examined in 5% potassium hydroxide (KOH),
Melzer’s reagent (IKI) and Cotton Blue (CB). Measurements and drawings were mainly
made in CB. A drawing tube was used when making line drawings. Iranian specimens
are maintained in Ghobad-Nejhad ref. herb. and some of dublicates are deposited in
H. Material from the Russian Caucasus and NE Turkey are kept in GB. The collecting
localities in NW Iran are mapped in Fig. 1 and the assigned numbers are explained
in Table 1. These numbers are used when examined specimens are listed, to avoid
redundancy. The author names are abbreviated as MG, NH, and HK in the text and/or
beside collection numbers. The nomenclature of species follows Cortbase version 2.02
(accessed 2007).
272 ... Ghobad-Nejhad, Hallenberg & Kotiranta
New records
*e 1. Acanthophysellum minor (Pilat) Sheng H. Wu, Boidin & C.Y.Chien _ Fi. 2
=Aleurodiscus cerussatus var. minor Pilat ;
=Acanthophysium minor (Pilat) Telleria
Aleurodiscus cerussatus var. minor (Pilat 1926) was raised to species rank
in Acanthophysium by Telleria (1991), and subsequently transferred to
Acanthophysellum by Wu et al. (2000). Telleria & Melo (1995) stated that
Acanthophysium cerrusatum differs from A. minor by spore size: 10-14 x 7-
9 um in the former and 7-10 x 5-7 um in the latter. Spores in the Iranian
specimens measure 6.5—9.5 x 3.5-6 um.
On dead wood of both angio- and gymnosperms. Most of the known
distribution for the species is given under Acanthophysellum cerussatum (Bres.)
Parmasto in a wide sense (=Aleurodiscus cerussatus (Bres.) Hohn. & Litsch.)
which is also known in Southern Hemisphere (see Hjortstam & Ryvarden 2007a,
b). According to Telleria & Melo (1995), A. minor is common on branches of
different hardwoods in dry and sunny places in Iberian Peninsula.
SPECIMENS EXAMINED — Iran. Loc (3), on fallen twig under Juniperus sp. and Paliurus
spina-christi, 01.X.2006 MG 363; on litter under Juniperus sp., MG 366. Croatia. Mt.
Vipera, on Pinus nigra, IV.1940, K.H. Rechinger & Litschauer, no. 1538 (H).
Fig. 2. Acanthophysellum minor. a) Section of hymenium with gloeocystidia (a,) and acanthophyses
(a,), b) basidia, c) dendrohyphidia, d) basidiospores. MG 366. Scale bar=10 um.
New corticioids from the Iranian Caucasus ... 273
Caer OY an
d
a) Basidia, b) young cystidium, c) mature cystidium, d) basidiospores. MG 383.
r=
=
=|
Fig. 3. Amylostereum laevigatum.
e 2. Amylostereum laevigatum (Fr.) Boidin FIG. 3
A. laevigatum is closely related to A. ferreum (Berk. & M.A. Curtis) Boidin &
Lanq. (see e.g. Tabata et al. 2000, Slippers et al. 2002), which, however, is known
only from S. America.
The species was earlier reported in the NW Caucasus by Mukhamedshin
(1992) on Taxus in Fagus-Abies belt (1000-1350 m). In Iran it was collected on
bark of a conifer, fallen in a mixed open forest, and in Russian Caucasus it was
found on log of Taxus in mixed deciduous forest of colchic type with evergreen
understory.
274 ... Ghobad-Nejhad, Hallenberg & Kotiranta
The species has been recorded from conifers in Northern Hemisphere. In Japan,
it was reported as a symbiont occurring in mycangia of a female wasp and
attacks Abies firma, A. homolepis, Cryptomeria japonica, and Picea jezoensis,
leading to the discoloration of the wood (Tabata & Abe 1999). In Switzerland it
is associated with stem canker of Taxus baccata (Hassler et al. 2004).
SPECIMENS EXAMINED — Iran. Loc. (4), on conifer, 02.X.2006 MG 383. Russia.
Krasnodar, 2 km NE Chosta, on Taxus, 09.VIII.1996 NH 12967.
Fig. 4. Dendrocorticium polygonioides.
a) Subicular hyphae, b) basidia, c) dendrohyphidia, d) basidiospores. MG 504.
Scale bar=10 um.
*e 3. Dendrocorticium polygonioides (P. Karst.) M.J. Larsen & Gilb.
=Corticium polygonioides P. Karst PIiGo4, PHOTOsm
In the specimen from Iran, dendrohyphidia are less branched and spores afe
shorter (6.1 x 4.1 um), compared to Larsen & Gilbertson (1977) and Maekawa —
(1994). This may refer to the young stage of the fruitbody.
On different hardwoods. Africa and Europe. Asia: Japan on decaying branch
of Fagus crenata (Maekawa 1994), Russia, Nizhny Novogrod region, on Quercus
New corticioids from the Iranian Caucasus ... 275
Photos. 1-5. Photographs of basidiocarps taken in the field. Photo. 1. Dendrocorticium polygonioides,
MG 504. Photo. 2. Megalocystidium luridum, MG 455. Photo. 3. Scytinostroma aluta, MG 369.
Photo. 4. Hyphoderma nemorale, MG 480.
(Spirin 2002, mentioned to be one of the characteristic species of virgin oak-
forest in that region), Iran.
SPECIMEN EXAMINED — Iran. Loc (10), on decorticated fallen branch of Quercus sp.,
decay class 3, 10.X.2006 MG 504.
°4. Hyphoderma litschaueri (Burt) J. Erikss. & A. Strid Fic. 5
The earlier report of H. litschaueri from Iran by Hallenberg (1981) was later
shown to be Hyphoderma nemorale (see below). Mukhamedshin (1992) reports
H. litschaueri from NW Caucasus on Betula in subalpine belt (1650-2200 m).
However, we could not confirm the identification.
In NW Iran it was collected on fallen log of Carpinus betulus, and in Russian
Caucasus on branch of Quercus in Quercus forest with Fagus sylvatica, Prunus
and Rhododendron.
On decayed wood of different angiosperms. North America, Spain (GBIF:
www.gbif.org), Norway (Hansen & Knudsen 1997), Sweden (Toresson 1994),
Carpathian Mountains (Eriksson & Ryvarden 1975), Italy (Mayrhofer et al.
2001), Russia, South America and Australia (Hjortstam & Ryvarden 2007a),
Iran.
276 ... Ghobad-Nejhad, Hallenberg & Kotiranta
a
a
Fig. 5. Hyphoderma litschaueri.
a) Basidia, b) cystidia, c) basidiospores. MG 462. Scale bar=10 um.
SPECIMENS EXAMINED — Iran. Loc. (8), on Carpinus betulus, 05.X.2006 MG 462.
Russia. Krasnodar, Lazarevskaja , alt. 410 m., N43°41°: E39°42’ on Quercus, 11.IX.1996
NH 12999 and NH 13006.
*e5. Hyphodontia erastii Saaren. & Kotir. Fic. 6
H. erastii belongs to Hyphodontia sambuci (Pers.) J. Erikss. group and is
distinguished by having fewer subulate cystidia, smaller capitate cystidia and
narrower, subcylindrical spores (Kotiranta & Saarenoksa 2000).
Four Iranian specimens were collected on fallen twigs of Carpinus betulus —
and on decayed twig of deciduous tree.
So far known on: Angelica sylvestris, Arctium tomentosum, Caragana sp.,
Carpinus betulus, Filipendula ulmaria, Phragmites, Sambucus kamtschatica,
and S. racemosa, mostly in semi-natural forests. Finland, Russia (Kotiranta &
New corticioids from the Iranian Caucasus ... 277
ti
Fig. 6. Hyphodontia erastii.
a) Hyphae of subiculum, b) Basidia , c) subulate cystidium,
d) capitate cystidium, e) basidiospores. MG 430.
Saarenoksa 2000), the Netherlands (Heller & Keizer 2004), regarded as critically
endangered in Czech Republic (Holec & Beran 2006), Iran.
SPECIMENS EXAMINED — Iran. Loc. (1), on deciduous twig, 29.1X.2006 MG 328: Loc.
(6), on Carpinus betulus, 04.X.2006 MG 430; Loc. (8), on Carpinus betulus, 05.X.2006
MG 445, 465.
°6. Megalocystidium luridum (Bres.) Jiilich Fic. 7, PHOTO. 2
=Gloeocystidiellum luridum (Bres.) Boidin
Known from Azerbaijan on branch of Pterocarya pterocarpa (Parmasto 1965,
denoted: exemplar non typicum). ‘The single specimen from Iran was found on
moderately decayed, decorticated standing stump of Carpinus betulus.
278 ... Ghobad-Nejhad, Hallenberg & Kotiranta
Mostly on angiosperms. Europe, Morocco (Malen¢on 1982), China (Dai et
al. 2004), Azerbaijan, Iran.
SPECIMENS EXAMINED — Iran. Loc (8), on Carpinus betulus, 05.X.2006 MG 455.
Fig. 7. Megalocystidium luridum.
a) Basidia, b) gloeocystidia, c) basidiospores. MG 455.
Scale bar=10 tm.
New corticioids from the Iranian Caucasus ... 279
*e7. Melzericium bourdotii Jiilich Hig: s
Julich (1976) distinguished M. bourdotii from Melzericium udicola (Bourdot)
Hauerslev essentially based on spore shape, which is “distinctly constricted in
the middle (like a dumb-bell)” in the former. Still, the taxon with bean- or
kidney-shaped spores is called M. udicola in most literature. A re-examination
of both types combined with DNA analysis is needed to solve if two taxa are
involved in practice.
The specimen from Iran was collected on fallen twig of Juniperus
communis.
Only sparse data is available for this species. Canada, Ontario, on Populus?
(de Vries 2004), France on Salix caprea (type), Norway on hard moist fallen
Fig. 8. Melzericium bourdotit.
a) Hyphae, b) basidia, c) basidiospores. MG 395A.
280 ... Ghobad-Nejhad, Hallenberg & Kotiranta
d
Fig. 9. Peniophora laeta.
a) Basidia, b) gloeocystidium, c) encrusted cystidia, d) basidiospores. MG 327.
New corticioids from the Iranian Caucasus ... 281
trunk of Alnus (Strid 1975, Ryvarden et al. 2003), Sweden (Hauerslev 1974).
The only known collection on conifer is from Iran, on Juniperus communis.
SPECIMENS EXAMINED — Iran. Loc. (4), on Juniperus communis, 02.X.2006 MG 395A.
*e8. Peniophora laeta (Fr.) Donk FIG. 9
Reminiscent of Peniophora incarnata (Pers.) P. Karst. which is found on cut
surfaces of wood. P. incarnata grows on different hosts, whereas P laeta inhabits
primarily corticated branches of Carpinus betulus where it grows on primary
cortex and pushes away the secondary layers of bark with hyphal pegs (e.g.
Eriksson et al. 1978).
Zugmaier et al. (1994) studied the mycoparastism of Peniophora laeta by
Tremella mesenterica Retz., growing on Carpinus betulus. In Iranian material,
at least in one occasion (no. 413A) the fruitbodies of T. mesenterica were close
to P. laeta.
In NW Iran P. laeta commonly inhabits corticated fallen or live twigs and
branches of Carpinus betulus. Found in Russian Caucasus on Carpinus in mixed
riverside forest, and in NE Turkey in mixed forest on unidentified deciduous
branch.
Mostly on corticated branches of Carpinus betulus in Europe. In N America
on Amelanchier pallida and A. utahensis (Ginns & Lefebvre 1993). Unterseher
& Tal (2006) recorded it on strongly decayed wood of Tilia cordata in Germany.
Apparently its easternmost known locality is in Iran.
SPECIMENS EXAMINED — Iran. Loc. (1), on hardwood twig, 29.1X.2006 MG 327 and
on Carpinus betulus, MG 337; Loc. (5), on Carpinus betulus, 03.X.2006 MG 413A; Loc.
(11), on Carpinus betulus, 29.11X.2005 MG 119. Russia. Krasnodar, Lazarevskaja, on
Carpinus, 19.1X.1996 NH 13150. Turkey. Trabzon, Sumela monastery, 1000-1100 m, on
hardwood branch, 02-04.X.1989 NH 11381.
¢ 9. Scytinostroma aluta Lanq. Fig. 10, PHOTO 3
Hallenberg (1985) discussed similar species and their distinguishing features.
S. aluta was earlier recorded from Stavropol, Russian Caucasus by
Bondartseva & Parmasto (1986). In NW Iran it was collected on dead but
still attached, corticated branches of Paliurus spina-christi, and on fallen twig
of Lonicera sp. In both cases the substrate was white-rotted. It has also been
sampled in NE Turkey on Sambucus nigra branch in a mixed Picea orientalis
forest with Fraxinus and Alnus barbata.
Submediterranean species (Hallenberg 1985) on different hardwoods.
Bondartseva & Parmasto (1986) give more extended account of its hosts and
distribution.
SPECIMENS EXAMINED — Iran. Loc. (3), on Paliurus spina-christi, 01.X.2006 MG 369;
Loc. (6), on Lonicera sp., 04.X.2006 MG 434. Turkey. Trabzon, Sumela monastery, 1000-
1100 m., on Sambucus, 02—04.X.1989 NH 11368.
282 ... Ghobad-Nejhad, Hallenberg & Kotiranta
10 um
C
Fig. 10. Scytinostroma aluta.
a) Basidia, b) cystidium, c) dendrohyphidia, d) basidiospores. MG 434.
*e 10. Tubulicrinis cinctus G. Cunn. FIG. 11
=Tubulicrinis cinctoides Hjortstam
The cystidia in Iranian specimen are frequently coiled with the hyphae of
another fungus (Fig. 12c). These may belong to Colacogloea bispora (Hauerslev)
Oberw. & R. Bauer. However basidia with only one sterigma were visible (not
shown in Fig. 11), and the other sterigma is apparently behind the cystidia.
Moreover the shape of the spores does not fully agree with C. bispora, and only
a few germinated spores were seen, but no really repetitive spores. Therefore we
are not fully sure of its identity. Only few encrustations were seen around the
neck of T: cinctus cystidia.
New corticioids from the Iranian Caucasus ... 283
Fig. 11. Tubulicrinis cinctus.
a) Hyphae in subiculum, b) basidia,
c) cystidia frequently coiled with the hyphae of unknown fungus (c,), d) basidiospores. MG 472A.
284 ... Ghobad-Nejhad, Hallenberg & Kotiranta
In Iran, on stump of Taxus baccata, accompanied with Radulomyces rickii.
Russian Caucasus, on a fallen log of Abies on northern slope of an Abies forest.
On dead wood of conifers viz. Abies, Dacrydium cupressinum, Picea, and
Taxus baccata. N Europe, Russia, Turkey, Iran, China (Maekawa et al. 2002),
New Zealand.
SPECIMENS EXAMINED — Iran. Loc. (9), on Taxus baccata, 06.X.2006 MG 472A. Russia.
Krasnodar, Mostovskoj, 1700 m, on Abies, 11.1X.1991 NH 12182.
Fig. 12. Vuilleminia coryli.
a) Basidia, b) dendrohyphidia, c) cystidium, d) basidiospores. MG 319.
New corticioids from the Iranian Caucasus ... 285
¢ 11. Vuilleminia coryli Boidin, Lanq. & Gilles Fic. 12
Vuilleminia coryli comes close to V. cystidiata Parmasto but the latter has
subulate cystidia (obtuse in V. coryli) and smaller spores.
Reported from NW Caucasus by Mukhamedshin (1992) on Corylus in
Quercus belt (650-750 m), Fagus belt (750-1000 m), and Fagus-Abies belt
(1000-1350 m).
In Iran on decorticated snag of Crataegus sp. This substrate seems unusual
for V. coryli, since the association with hosts from Rosaceae, and Crataegus in
particular, has used to be a distinguishing feature for V. cystidiata. In Russian
Caucasus, as well as NE Turkey, V. coryli was consistently collected on Corylus
in a variety of luxuriant forests.
Usually on Corylus. England (Legon et al. 2005, also single collections on
Acer campestre, Salix caprea and Sambucus nigra), France (Boidin et al. 1989,
once on Acer campestre), Basque (Salcedo 1994), Luxemburg (Tholl et al. 2000),
N Europe (Hansen & Knudsen 1997), Russia, Turkey, Iran.
SPECIMENS EXAMINED — Iran. Loc. (1), on Crataegus sp., 29.IX.2006 MG 319. Russia.
Krasnodar, Mostovskoj 1700 m, 11.1X.1991 NH 12164; Lazarevskaja , Baranovka, alt. 60
m., N43°42°: E39°41’, 08.VIII.1996 NH 12911; Krasnodar, Lazarevskaja , 19.IX.1996
NH 13142; Adler, Estosadok. alt. 240 m, 15.IX.1996 NH 13091. Turkey. Trabzon,
vicinity of Malavaci. alt. 1200 m, N40°52’: E39°37’, 26.1X.1996 Hallenberg 13262. (all
on Corylus).
Additional finds of species rarely collected in Iran
Twelve species collected previously only once or twice in Iran are listed, and
their occurrence in the Caucasus is summarized.
1. Athelia arachnoidea (Berk.) Jiilich FIG. 13
Although A. arachnoidea is a quite commen species, the only earlier report is
the two collections made by Hallenberg (1981) on fallen branches in N Iran.
In NW Iran five samples (all fertile) were collected on fallen corticated
branches and twigs or on fallen bark of Quercus and Paliurus spina-christi.
Spore shape in one collection is unusual (Fig. 14) and their size is small, 6.5 x
5.5 um. One fruitbody (no. MG 306) was accompanied with Stereum hirsutum,
and one collection (no. MG 373) parasitized the lichen Pleurosticta acetabulum.
No data on the occurrence of its anamorph Fibulorhizoctonia carotae (Rader)
G.C. Adams & Kropp in Iran was found.
In NE Turkey collected on Picea log, in Fagus, Abies, Picea forest.
Mostly on angiosperms, rarely recorded on gymnosperms. Distributed in all
continents, but reports from Asia are few, not yet known in China and Japan
(cfr. Dai et al. 2004 and Maekawa 1993, respectively).
286 ... Ghobad-Nejhad, Hallenberg & Kotiranta
ae j dni
D000
Fig. 13. Athelia arachnoidea.
a) Hyphae in subiculum, b) a section of hymenium, c) basidia, d) basidiospores. MG 306.
Scale bar=10 um.
SPECIMENS EXAMINED — Iran. Loc. (1), on Quercus, 29.[X.2006 MG 306; Loc. (3), on
Paliurus spina-christi, MG 373, on felled bark, MG 375; Loc. (4), on Quercus, MG 376;
Loc. (10), on Quercus, MG 491. Turkey. Cilekli Agacli, 1400 m, on Picea, 10.X.1989 NH
Blois:
2. Ceraceomyces serpens (Tode) Ginns
=Ceraceomerulius serpens (Tode ) J. Erikss. & Ryvarden
Earlier records were made by Hallenberg (1981) on a fallen branch of Fagus
orientalis in N Iran, by Mukhamedshin (1992) in Quercus belt (650-750 m)
and Fagus belt (750-1000 m) on Sambucus and Corylus in NW Caucasus.
Gvritishvili et al. (2007) mention it for Borjomi, Georgia. New collections were
made on decorticated fallen branches of Juniperus communis. j
Found in many parts of the world, on both angio- and gymnosperms.
SPECIMENS EXAMINED — Iran. Loc. (4), on Juniperus communis, MG 383a, 395.
New corticioids from the Iranian Caucasus ... 287
3. Corticium roseum Pers.
=Laeticorticium roseum (Pers.) Donk
Earlier reports in N Iran: on fallen logs of Fagus and Quercus (Hallenberg
1981). In the Caucasus: Gardabani and Abkhasia, Georgia (Gvritishvili et al.
2007); NW Caucasus on Salix, Populus, and Fagus in all four vegetation belts
(see Mukhamedshin 1992).
Collections in NW Iran were made on decorticated and corticated, dead, but
still attached branches and twigs of Carpinus betulus and on cut surface of an
unidentified dead angiosperm. In Russian Caucasus on a still attached branch
of Populus and log of Alnus in mixed forests.
More often on hardwood. Found in all continents except in S America.
SPECIMENS EXAMINED — Iran. Loc. (8), on Carpinus betulus, MG 448; Loc. (9), on
Carpinus betulus, MG 471; Loc (10), on angiosperm, MG 518. Russia. Krasnodar,
Mostovskoj, alt. 1000 m, on Populus, 07.1X.1991 NH 12037; Adler, N43°39’: E40°27’
alt. 930 m, on Alnus, 21.1X.1996 NH 13238; Estosadok. alt. 240 m, on Alnus, 15.1X.1996
NH 13086.
4. Hyphoderma nemorale K.H. Larss. Fic. 14, PHOTO. 4
Denoted for Iran by Larsson (1998) on Fagus orientalis (Hallenberg 1981: sub
Hyphoderma cremeoalbum (Hohn. & Litsch.) Jiilich and H. litschaueri). In
the present study 10 collections in Arasbaran were made on Carpinus betulus
and Quercus sp., mostly on dead, fallen, decorticated branches and twigs,
which sometimes were considerably decayed. On some branches were also
Hyphoderma setigerum s.1., Irpex lacteus, and Skeletocutis nivea.
Often found on small corticated branches and twigs of different hardwoods
in boreal and nemoral zones (Larsson 1998). The occurrences here on mostly
decorticated branches point towards Hyphoderma incrustatum K.H. Larss.
(Larsson 1998), but microscopically they fit H. nemorale.
No data about the species in other parts of the Caucasus is available.
SPECIMENS EXAMINED — Iran. Loc (1), on Carpinus betulus, 29.[X.2006 MG 339; on
angiosperm, MG 340a; Loc. (5), on C. betulus, 03.X.2006 MG 417; on Quercus, MG 421;
Loc. (8), on fallen twig of C. betulus, 05.X.2006 MG 444; on C. betulus, MG 449, 456;
on C. betulus, MG 454; Loc. (9), on angiosperm, 06.X.2006 MG 480 and on Quercus,
MG 489.
5. Hyphodontia subalutacea (P. Karst.) J. Erikss.
Previously reported from N Iran by Hallenberg (1981) on fallen, white-rotted
branches of Fagus orientalis. From NW Caucasus by Mukhamedshin (1992) on
Pinus in Abies belt (1350-1650 m) and Subalpine belt (1650-2200 m).
The majority of the reported collections were made on conifer branches.
In Iran on dead, decorticated hanging branch of Juniperus communis , dead,
decayed but still attached burnt branch of Cornus mas, and on brown-rotten
288 ... Ghobad-Nejhad, Hallenberg & Kotiranta
Fig. 14. Hyphoderma nemorale.
a) Hyphae in subiculum, b) basidia, c) hymenial cystidia, d) enclosed cystidia,
e) basidiospores. MG 417. Scale bar=10 um.
fallen branch of Quercus. In Russia on Abies in Abies forest and on Pinus in
Pinus-Quercus forest. In Turkey on Picea in a pure Picea orientalis stand.
On both angio- and gymnosperms with worldwide distribution.
SPECIMENS EXAMINED — Iran. Loc. (1), on fallen decorticated branch of Quercus; Loc.
(4), on Juniperus communis, 02.X.2006 MG 384; Loc. (9), on Cornus mas, 06.X.2006 MG
473, Russia. Krasnodar, Mostovskoj, 1700 m, on Abies, 11.IX.1991 NH 12186; 1000 m,
on Pinus, 13.[X.1991 NH 12250. Turkey. Trabzon, 1600 m, on Picea, 02—04.X.1989 NH
W299 L293;
6. Peniophora nuda (Fr.) Bres.
Reported by Hallenberg (1981) on fallen branches in N Iran. In the Caucasus -
Mukhamedshin (1992) on Fagus and Carpinus in all four vegetation belts (650-
1650 m) in NW Caucasus. It also occurs in Abkhasia Autonomous Republic,
Akhmeta, and Gardabani, Georgia (Gvritishvili et al. 2007).
New corticioids from the Iranian Caucasus ... 289
Our material is from hardwood branches in deciduous or mixed forests: NW
Iran on bark of fallen hardwood branch, with Stereum hirsutum, Russian
Caucasus on branch of Fagus in a riverside forest and on Carpinus branch in
mixed forest, Turkey on an unidentified hardwood branch in a mixed forest.
Known from all continents, preferably on angiosperms, a few times on
Juniperus (e.g. Ginns & Lefebvre 1993, Garcia-Manjon & Moreno 1981).
SPECIMENS EXAMINED — Iran. Loc. (1), on hardwood branch, 29.1X.2006 MG 304.
Russia. Krasnodar, Mostovskoj, on Fagus, 06.1X.1991 NH 11995; Umpyr, 1000 m,
on Carpinus, 07.1X.1991 NH 12003 (P. nuda s.1.). Turkey. Trabzon, 1000-1400 m, on
hardwood branch, 02—12.X.1989 NH 11591.
7. Peniophora violaceolivida (Sommerf.) Massee
Reported by Hallenberg (1981) on fallen branches and twigs in N Iran. On
Carpinus and Vaccinium arctostaphylos in all four vegetation belts (650-1650
m) in NW Caucasus (see Mukhamedshin 1992).
A species with a wide distribution, preferably on deciduous trees, seldom on
conifers (e.g. Ginns & Lefebvre 1993).
SPECIMEN EXAMINED — Iran. Loc. (2) on fallen stump of Populus sp., 30.1X.2006 MG
358.
8. Terana caerulea (Lam.) Kuntze
=Pulcherricium caeruleum (Lam.) Parmasto
In Iran, earlier found only once on a fallen log of Quercus castaneifolia
(Hallenberg 1981). In Abkhasia Autonomous Republic, Georgia (Gvritishvili et
al. 2007), Armenia (Melik-Hacatryan 1971), on Corylus and Quercus in Ordu,
Turkey (Dogan et al. 2005).
Newly collected in NW Iran on bark of moderately decayed, dead hanging
branches of Carpinus betulus and Cornus mas.
More or less with a wide distribution range (apparently not reported in S
America), in sites with warm and humid (micro) climate. Always found on
hardwoods.
SPECIMENS EXAMINED — Iran. Loc. (9), on Carpinus betulus, 06.X.2006 MG 469 and on
Cornus mas, MG 474.
9. Tubulicrinis incrassatus Hallenb.
Described from N Iran by Hallenberg (1978) on fallen branches of deciduous
tree in a closed forest consisting of Parrotia persica, Carpinus betulus, Quercus
castaneifolia and Acer. In NW Iran on fallen branch of Juniperus with advanced
brown rot, in a dry and rather dense Juniperus shrubland.
In Iran, T. incrassatus is known from different types of vegetation, on
hardwood and on conifer. According to Hjortstam et al. (1988), it was collected
290 ... Ghobad-Nejhad, Hallenberg & Kotiranta
in the southern part of Italy. Hjortstam & Ryvarden (2007b) report Tubulicrinis
cf. incrassatus from Venezuela.
SPECIMEN EXAMINED — Iran. Loc. (3), on Juniperus, 01.X.2006 MG 365.
10. Xenasma pruinosum (Pat.) Donk BiG:15
Two collections were made by Hallenberg (1981) in N Iran on a fallen branch
of Parrotia persica and an unknown tree.
Known from all continents but Australia. Often on decayed deciduous
trees.
SPECIMEN EXAMINED — Iran. Loc. (4), on decayed stump of hardwood, with
Peniophorella praetermissa, 02.X.2006 MG 380a.
ay ctacl «niet
q
OO
Fig. 15. Xenasma pruinosum.
a) Basidia, b) cystidiole with apical protuberances, c) cystidium, d) basidiospores. MG 380a.
Scale bar=10 um.
New corticioids from the Iranian Caucasus ... 291
Remarks
About 170 species of corticioids are known from Iran. Indeed this figure comes
out of few inventories in the northern forest belt of the country, and therefore is
far from complete. Any conclusion on the species composition of the country
will be premature. Even the species mentioned in the second part of this paper
may proven to be more abundant if searched for. In fact many of them are
globally widely distributed. Representatives of the two genera Aleurodiscus
s.l. and Amylostereum are reported for the first time from Iran. The known
distribution of Tubulicrinis incrassatus is apparently confined to Iran and Italy.
Acknowledgments
MG is grateful to CIMO and Societas pro Fauna et Flora Fennica for financial support.
Peter Roberts (Kew) and Leif Ryvarden (Oslo) have kindly reviewed the manuscript and
are warmly thanked. Mohammad Sohrabi (Helsinki) assisted actively in the field work
in Iran.
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Volume 105, pp. 295-300 July-September 2008
Entoloma reinwaldii,
a rare species new to Croatia
ARMIN MESIC’ & ZDENKO TKALCEC
amesic@irb.hr ztkalcec@irb.hr
Laboratory of Biocoenotic Research, Ruder Boskovic Institute
Bijenicka cesta 54, HR-10000 Zagreb, Croatia
Abstract — Entoloma reinwaldii is recorded as new to Croatian mycobiota and
described. The description is accompanied by black and white photographs of fresh
basidiocarps and microscopic characters.
Key words — Entolomataceae, Agaricales, taxonomy, biodiversity, biogeography
Introduction
During the research of rockrose garigue (Cistus spp.) mycobiota in
Mediterranean part of Croatia in 2000, we found two collections of beautiful
pink-colored species of Entoloma (Fr.) P. Kamm. After microscopic examination
and consultation of the literature it became clear that our collections belonged
to a then undescribed species of subgenus Leptonia (Fr.) Noordel., section
Cyanula (Romagn.) Noordel. Soon after that, Noordeloos & Hausknecht (2001)
described this species as new, under the name Entoloma reinwaldit, from a
similar Mediterranean habitat in Italy.
Materials and methods
Our study of Entoloma reinwaldii is based on two collections consisting of
nine basidiocarps. The photograph of basidiocarps was taken in the field.
The description of macroscopic characters is based on observations of fresh
basidiocarps. Basidiocarps were preserved by drying. Microscopic features were
observed by a light microscope (brightfield) with magnification up to 1500 x
and photographed with a digital camera. The description and photographs of
microscopic characters were made from rehydrated dried specimens mounted
* Corresponding author
296 ... Mesié & Tkaléec
in 5% potassium hydroxide (KOH) solution. Spore measurements were made
from the mounts of lamellae and based on calibrated digital photographs.
One mature basidiocarp was selected from each collection and 50 randomly
selected spores (100 spores in total) were measured without the apiculus.
Spore measurements (length, width) are given as: (min.) stat. min.—av.-stat.
max. (max), where “min.” = minimum (lowest measured value), “stat. min” =
statistical minimum (arithmetic average minus two times standard deviation),
“av.” = arithmetic average, “stat. max.’ = statistical maximum (arithmetic
average plus two times standard deviation), “max.” = maximum (highest
measured value). The range of arithmetic averages (av.) of spore measurements
of each particular collection is also given. Standard deviation (SD) of spore
length and width is given as: min.-total-max, where “min.” = collection with
lower SD value, “total” = SD value of all 100 measured spores, and “max” =
collection with higher SD value. The length/width ratio of spores is given as the
“Q” value (min.—av.-max.), and the range of arithmetic averages of “Q” value
(Q av.) of each particular collection is also given. Both Croatian collections
with accompanied data are deposited at the Croatian National Fungarium in
Zagreb (CNF).
Taxonomic description
Entoloma reinwaldii Noordel. & Hauskn.,
Boll. Gruppo Micol. G. Bresadola, n.s. 43(3): 26, 2001. FIGS 1-9
Pit—Eus 13-22 mm broad, subhemisphaerical to truncately conical at first,
later convex to plano-convex, often with applanate or depressed center,
hygrophanous, pink, lilac pink, orangish pink or brownish pink when moist,
pale pink on drying, more intensely colored at center, translucently striate
up to 3/4 of radius, minutely squamulose, also somewhat radially fibrillose
near margin, dry. LAMELLAE adnexed to emarginate, moderately distant to
moderately crowded, whitish to pinkish (sometimes with lilac tinge) at first,
then dirty pink, with entire, concolorous edge. STIPE 35-60 x 1.5-2.5 mm,
subcylindrical, slightly broadened at apex, dirty pink, minutely pubescent to
pruinose under a hand-lens (especially at apex), lower part often glabrous,
white tomentose at base, dry, fistulose. CONTEXT whitish to pinkish. SMELL
weak, pleasant. Taste mild, somewhat fruity.
Spores (9.6-)10.0-11.3-12.5(-12.7) x (6.7-)6.8-7.8-8.9(-9.1) ym, av. 11.2-
11.3 x 7.8-7.9 um; SD = 0.64-0.64-0.64 x 0.47-0.51-0.55, Q = 1.22-1.44-1.68,
Q av. = 1.43-1.45, heterodiametrical, with (5—)6-8(-9) regular to irregular
angles in side view, dirty yellowish with some pink reflections in KOH (pink
color more visible in water), thin-walled. Basip1A 25-36 x 9-13 um, clavate to
subcylindrical, 4-spored, thin-walled, clampless. LAMELLAREDGEsterile towards
the margin of the pileus, heterogeneous towards the stipe. CHEILOCYSTIDIA
Entoloma reinwaldii in Croatia ... 297
Figs 1-3. Entoloma reinwaldii. 1. Basidiocarps. 2. Spores. 3. Basidia.
Bars: 1 = 10 mm; 2 & 3 = 10 um.
usually septate, thin-walled, hyaline, terminal elements clavate to subcylindrical,
17-61 x 6-14 um. PLEUROCYSTIDIA absent. HYMENOPHORAL TRAMA regular,
composed of 1.5-23 um broad, thin-walled, hyaline hyphae. PILEIPELLIs a
cutis (except at disc) composed of hyaline to brownish, thin-walled hyphae,
with scattered bundles of repent to ascending, clavate to subcylindrical, 7-20
um broad terminal elements; a hymeniderm to trichoderm at disc, composed
of inflated elements, 23-50 x 12-28 um; pigment intracellular. REFRACTIVE
HYPHAE abundant in pileitrama, also present in hymenopodium, 2.5-8(-10)
um broad, yellowish, thin-walled. ST1PITIPELLIS a cutis composed of thin-
walled, hyaline hyphae, with scattered bundles of ascending, + cylindrical
elements (especially at apex of stipe). CLAMP CONNECTIONS absent.
298 ... Mesié & Tkaléec
Figs 4-9. Entoloma reinwaldii. 4. Cheilocystidia. 5. Stipitipellis. 6. Pileipellis (cutis) and
pileitrama with refractive hyphae. 7. Bundles of repent terminal elements in pileipellis. 8.
Bundles of ascending terminal elements in pileipellis. 9. Pileipellis at disc (hymeniderm).
Bars = 10 um.
on
Entoloma reinwaldii in Croatia ... 299
HasitaT: Garigue with Cistus monspeliensis, C. salviifolius, C. incanus, Erica
arborea, Juniperus oxycedrus subsp. macrocarpa, and Phillyrea sp., on soil.
COLLECTIONS EXAMINED: CROATIA, Istria, Premantura peninsula (near the town of
Pula), 44°46°07"N, 13°55°04”E, alt. 25 m, 2 December 2000, leg. Z. Tkaléec & A. MeSi¢
(CNF 1/2111, 1/2117).
Discussion
Entoloma reinwaldii is characterized by pink basidiocarps, collybioid habit,
minutely squamulose, hygrophanous, and translucently striate pileus, pileipellis
a trichoderm to hymeniderm at center and a cutis with scattered bundles
of repent hyphae elsewhere, absence of clamp connections, and clavate to
cylindrical cheilocystidia. In the section Cyanula, nine more European species
with pink or distinctly pink tinged pileus are known. Entoloma catalaunicum
(Singer) Noordel., E. queletii (Boud.) Noordel., E. roseotinctum Noordel. &
Liiv, E. roseum (Longyear) Hesler, and E. rufocarneum (Berk.) Noordel. can be
differentiated from E. reinwaldii by their not hygrophanous and not (or hardly)
translucently striate pileus. Entoloma ianthinum (Romagn. & J. Favre) Noordel.
differs by complete absence of cheilocystidia, E. ursulae Noordel. et al. differs
by dark blue stipe, E. callirhodon Hauskn. & Noordel. differs by darker (reddish
lilaceous), fimbriate lamellar edge, and E. cyanulum var. roseolum Noordel. &
Sullock-Enzlin differs by somewhat larger spores, papillate pileus, and smaller
basidiocarps. Descriptions and illustrations of these species can be found in
Noordeloos (1992, 2004).
Entoloma reinwaldii is a rare species. According to our knowledge, hitherto it
has been found in four European countries: Italy (Noordeloos & Hausknecht
2001), France (Lejeune 2001), Spain (Vila & Caballero 2007), and Croatia.
In each of these countries, the species is known from only one locality. In
Italy, it was found on 19 and 21 November 1997 (two records), in province of
Foggia, Mattinata, Tratturita, in Mediterranean macchia of Pistacia lentiscus,
Cistus monspeliensis, Quercus coccifera, and Q. ilex. In France, it was found
on 10 September 2000, in Robertsau forest at the periphery of Strasbourg, in
a forest of Populus sp., Acer sp., and Corylus sp. In Spain, it was found on 7
October 2006, in Serra d’Heures, Sant Hilari Sacalm (Girona), in a forest of
Corylus avellana, Quercus pubescens, Fagus sylvatica and Fraxinus sp. It can
be concluded that Entoloma reinwaldii lives in different types of habitats with
wood or shrub vegetation, in Mediterranean and continental climates.
Our collections show some differences compared to collections described in the
literature. (i) Lilac tinges in the pileus, which are present in some basidiocarps
of our collections, are not mentioned by any other author. Moreover, in his
dichotomous key Noordeloos (2004: 814, 815) emphasized that the pileus of
300 ... Me’i¢ & Tkaléec
Entoloma reinwaldii has no lilaceous tinge. (ii) In our collections lamellae are
adnexed to emarginate, while in the other descriptions lamellae are said to be
more variable and also can vary from adnate to (sub)decurrent. (iii) Spores
in the French collection (Lejeune 2001) are slightly shorter ([8.8-]10-11.2 x
8 um). (iv) Noordeloos & Hausknecht (2001) and Lejeune (2001) described
lamellar edge as completely sterile, while basidiocarps in our collections have
sterile (towards the margin of the pileus) to heterogeneous (towards the stipe)
lamellar edges. The collection of Vila & Caballero (2007) also shows variability
of lamellar edge, where cheilocystidia are abundant in some basidiocarps and
absent in others. (v) All authors described pileipellis at the center of pileus as
a trichoderm, while basidiocarps in our collections have a hymeniderm to
trichoderm with more inflated elements. (vi) Our collections have abundant
refractive hyphae in pileal trama (also present in the hymenopodium), which is
not mentioned by the other authors.
Acknowledgements
We are grateful to Pierre-Arthur Moreau (Lille, France) and Fernando Esteve-Raventos _
(Alcala de Henares, Spain) for their critical reviews of the manuscript.
Literature cited
Lejeune C. 2001. Une leptonie italienne, de création récente, en forét du Rhin: Entoloma reinwaldii
Noordel. & Hauskn. Bull. Soc. Mycol. Strasbourg 80: 9-11 + cover.
Noordeloos ME. 1992. Entoloma s.]. Fungi Europaei 5. Libreria editrice Giovanna Biella, Saronno.
Noordeloos ME. 2004. Entoloma s.l. Supplemento. Fungi Europaei 5a. Edizioni Candusso, Alassio.
Noordeloos ME, Hausknecht A. 2001 “2000”. Tre nuove Entolomataceae (Agaricales) dall’Italia.
Boll. Gruppo Micol. G. Bresadola, n.s. 43(3): 23-33.
Vila J, Caballero F. 2007. Entoloma nuevos o interesantes de la Peninsula Ibérica. Fungi non
delineati 38. Edizioni Candusso, Alassio.
MY COTA XON
Volume 105, pp. 301-312 July-September 2008
Studies in the genus Entoloma (Basidiomycota, Agaricales)
from the Kiklades (C. Aegean, Greece)
MACHIEL NOORDELOOS! & ELIAS POLEMIS?
‘noordeloos@nhn.leidenuniv.nl
National Herbarium of the Netherlands
P.O. Box 9514, 2300 RA Leiden, The Netherlands
* eliasp@ath.forthnet.gr
University of Patras, Department of Biology, Panepistimioupoli, GR-26500 Rion, Greece
Agricultural University of Athens, Laboratory of General & Agricultural Microbiology,
Tera Odos 75 GR-11855 Athens, Greece
Abstract—An intensive mycological inventory of the Greek Kiklades Islands yielded a
number of interesting records, two of them being new to science: Entoloma leuconitens
and E. alnicola. Descriptions are given of another eight taxa new to the Greek mycoflora,
among them the extremely rare Entoloma nigroviolaceum, E. griseopruinatum, and
E. griseorugulosum.
Key words—new species and records, agarics, Entolomataceae
Introduction
Reports on the macrofungi from the islands of the Aegean archipelagos are
scarce in literature (Petrak 1943a,b, Plank 1980). In the frame of an ongoing
inventory of the Greek macrofungi (Dimou et al., 2008; Zervakis et al., 2004),
the islands of Kiklades (C. Aegean, Greece) Andros, Naxos and Amorgos
have been thoroughly investigated. In the recent past, Andros has been
visited periodically by P. Lizon (1993-1997), and from 1995 until present by
E. Polemis, who is conducting a year-by-year inventory (mainly from September
to March). Some preliminary results of this work, including several records new
for Greece, were presented in national check-lists (Zervakis et al., 1998; 1999)
and during scientific conferences (Polemis et al., 2002; 2007). Recently, two
new species of Gymnopus were described from the island of Andros (Polemis
& Noordeloos, 2007).
The genus Entoloma has been studied by the first author on a worldwide
scale (Gates & Noordeloos 2007, Manimohan et al. 2006, Noordeloos 1981,
1987, 1992, 2004, 2006, 2007). However, data are still lacking on occurrence
302 ... Noordeloos & Polemis
and distribution of many species in Southern Europe, in particularly from
the Eastern Mediterranean region (Balkans, Greece). To date, the biodiversity
of this major agaricoid genus, is very basically known in Greece, as only a
limited number of taxa (less than 30 in total), if compared with other European
countries, are recorded. This paper presents two new species from the Kiklades,
and describes another five rare, as well as three more widespread and common
European taxa, that so far had not been recorded from Greece.
Taxonomic part
1. New taxa
Entoloma leuconitens Noordel. & Polemis, sp. nov. Fig. 1
MycoBAnk MB 512002
Pileus c 20 mm latus, conicus, haud hygrophanus, paulisper translucido-striatus, albus,
glaber, sericeus. Lamellae subliberae, roseae. Stipes 20 x 2 mm, albus, politus. Odore
saporeque rancidis. Sporae (9-)9.5-12(-15) x (6.5-)7.5-9 um, 6-8-angulatae. Acies
lamellarum sterilis. Cheiloystidia 15-45 x 7-15 um, utriformia. Pileipellis cutis hyphis
cylindraceis, 2-6(-8) um latis, pigmentis intracellulosis. Fibulae praesentes. Ad terram
acidam. Holotypus: Greece, Isl. Andros, Kiklades (C. Aegean), Torna, 29 Oct. 2002, E.
Polemis 02-A472 (L)
ETYMOLOGY—Ieucos = white; nitens = shining.
DeEscripTion—Pileus 20 mm, conical, with slightly inflexed, undulating
margin, not hygrophanous, faintly translucently striate at the extreme margin
only, white to creamy, ivory, glabrous with a lustrous appearance. Lamellae
moderately distant, adnate to almost free (adnexed), white then pinkish, with
concolourous edge. Stipe 20 x 4 mm, cylindrical, slightly broadened towards
base, curved, white to creamy, smooth, glabrous, polished to faintly fibrillose
striate, brittle. Context thin, brittle, white, smell and taste rancid.
Spores (9-)9.5-12(-15) x (6.5-)7.5-9 um, Q = 1.2-1.5(-1.8),
heterodiametrical, with 6-8 rather irregular angles. Basidia 30-40 x 11-13
um, 4-spored. Lamellae edge sterile. Cheilocystidia 15-45 x 7-15 um, mostly
utriform, to cylindrical-clavate. Pleurocystidia absent. Pileipellis a thin cutis
made up of cylindrical, 2-6(-8) um wide hyphae. Pigment very faint, probably
intracellular. Clamps present in hymenium, not seen elsewhere.
HasitatT: In acidic soil in mountain heath-land with Erica arborea, and
Pteridium aquilinum and with scattered trees like Fraxinus sp., Crataegus
monogyna and Pyrus amygdaliformis, on soil among P. aquilinum.
COLLECTION EXAMINED: Greece, Isl. Andros, Kiklades (C. Aegean), Torna, 29 Oct. 2002,
E. Polemis 02-A472 (L, holotype).
The white, non-hygrophanous, lustrous pileus, elongate, rather irregularly
shaped spores, and sterile lamella edge with utriform cheilocystidia are
distinctive for this species. Entoloma pallideradicatum Hauskn. & Noordel. is
New Entoloma species from Greece ... 303
CH
Fig. 1. Entoloma leuconitens. Spores, cheilocystidia (CH), and pileipellis (PP). Bar = 10 um.
close, but differs in slightly smaller spores, heterogeneous lamella edge, and
lack of clamp-connections.
Entoloma alnicola Noordel. & Polemis, sp. nov. Fig. 2.
MycoBank mb 512003
Pileus 13-20 mm latus, convexus demum applanatus, hygrophanus, translucido-striatus,
obscure griso-brunneus, pallescens, minute fibrillosus. Lamellae adnatae, roseae. Stipes
20-30 x 2 mm, griseo-brunneus, fibrilloso-striatus, ad apicem fibrillosus. Odore saporeque
haud notatis. Sporae (7.5-)8.5-10.5(-11.5) x 6.5-8 um, 5-6(-7) angulatae. Acies
lamellarum sterilis. Cheilocystidia 15-45 x 6.5-17 wm, clavata, cylindracea vel utriformia.
Pileipellis trichoderma hyphis inflatis ad 30 um latis, pigmentis intracellulosis. Fibulae
absentes. At terram in alnetum. Holotypus: Greece, Isl. Andros, Kiklades (C. Aegean),
Torna, 22 Sep. 2002, E. Polemis 02-A364 (L; isotype LGAM-AUA )
EtymMo.ocy: alnicola = associated with Alnus.
304 ... Noordeloos & Polemis
CH
A
Ae
ee
Ss My
-A
Fig. 2. Entoloma alnicola. Spores, cheilocystidia (CH), and pileipellis (PP). Bar = 10 um.
PP
DEscRIPTION—Pileus 13-20 mm, finally planocovex to aplanate, hygrophanous,
translucently striate, dark grey-brown at disk and striae, fading to sordid brown
towards the margin, which is finally straight and slightly undulating, minutely
fibrillose-squamulose. Lamellae moderately distant, adnate, white to pinkish,
with concolorous edge. Stipe up to 20-30 x 2 mm, cylindrical, greyish-beige at
apex, darker grey-brown at base, fibrillose striate, pruinose at apex. Smell and
taste not recorded.
Spores (7.5-)8.5-10.5(-11.5) x 65-8 um, Q = 1.2-1.4(-1.5),
heterodiametrical, with 5-6(-7) pronounced angles, relatively thick-walled.
Basidia (1)2-spored. Lamellae edge sterile. Cheilocystidia 15-45 x 6.5-17 um,
broadly clavate to cylindrical, utriform. Pileipellis a trichoderm with inflated,
clavate terminal elements, up to 30 um broad. Brilliant granules present.
Pigment brown, intracellular. Clamps absent.
New Entoloma species from Greece ... 305
HasiTaT: Saprotrophic on wet mossy soil with woody residues of Alnus
glutinosa along a stream with Platanus orientalis, and Alnus glutinosa.
COLLECTION EXAMINED: Greece, Isl. Andros, Kiklades (C. Aegean), Torna, 22 Sept.
2002, E. Polemis 02-A364 (L, holotype).
Entoloma alnicola belongs to subgenus Cyanula, stirps Longistriatum, close to
E. longistriatum (Peck) Noordel., from which it differs, however, by the rather
dark grey-brown tinges in pileus and stipe, a fibrillose-striate stipe surface,
the 2-spored basidia and the predominantly utriform or broadly clavate
cheilocystidia. Entoloma turci (Bres.) M.M. Moser, which is similarly grey-
brown, has a less distinct, non-translucently striate pileus, polished stipe, which
frequently turns reddish at the base, 4-spored basidia and differently shaped
cheilocystidia. Entoloma scabropelle Noordel. has a fertile lamella edge, and 4-
spored basidia.
2. New Greek records
Entoloma nigroviolaceum (P.D. Orton) Hesler, Persoonia 11(4): 471 (1967)
DESCRIPTION—Pileus 10-35 mm, hemispherical to convex, irregularly convex,
flat at centre, margin involute and often undulate, not translucently striate, not
hygrophanous, initially dark violaceous-black, then vinaceous-grey, at first,
innately fibrillose-tomentose to rugulose at centre, then entirely tomentose,
radially fibrillose to squamulose at centre. Lamellae moderately distant, adnate,
whitish to grey-bluish (lead-grey), then pinkish with concolorous edge. Stipe
30-50 x 2-6 mm, cylindrical to compressed, grey-blue to violaceous-grey,
fibrillose-striate, initially whitish tomentose towards the base, pruinose at apex,
becoming smooth with age. Context pale blue-violaceous, Smell absent, weak.
Spores (9-)9.5-13 x 7-9 um, Q = 1.2-1.5, irregularly 5-8 angled in side
view. Basidia (2)4-spored. Lamellae edge fertile. Cystidia absent. Pilepellis a
transition between a cutis at margin and a trichoderm towards the centre, made
up of inflated terminal elements up to 25 um broad, some apically or laterally
rostrate, pigment grey to violaceous-brown, intracellular. Clamps absent.
HABITAT: Mountain acidic heath land with Erica arborea, Genista acanthoclada,
Crataegus monogyna, and Pteridium aquilinum, on wet mossy soil.
COLLECTION EXAMINED. Greece, Isl. Andros, Kiklades (C. Aegean), Vourkoti, 8 Oct.
2002, E. Polemis 02-A414.
Entoloma nigroviolaceum is a striking species with dark violaceous-black
basidiocarps, and large, many-angled spores. It is a very rare species, known
from only few records in Northwestern Europe (Noordeloos 2004). In all cases
it has been found on fairly poor, acid peaty soil with mosses and Sphagnum. It
has never been recorded so far from Greece. Entoloma nigroviolaceum belongs
306 ... Noordeloos & Polemis
to a small group of closely related species, all of them being rare and indicative
of very special habitat conditions.
Entoloma griseopruinatum Noordel. & Cheype, Fungi Europaei, 5a: 876 (2004)
DESCRIPTION—Pileus 40-60 mm, planoconvex, finally somewhat depressed
at centre, with a small umbo and deflexed, straight to undulating margin, not
hygrophanous, not or slightly translucently striate, dark grey-brown, radially
innately fibrillose, with a greyish pruina left at the centre of pileus. Lamellae
moderately distant, adnate-emarginate, grey-beige with a pink tinge, with
concolourous or slightly paler edge. Stipe 40-60 x 7-15 mm, cylindrical to
somewhat compressed, base somewhat rooted, concolorous with pileus, with
white fibrillose covering, pruinose at apex, whitish and almost smooth at base.
Context thick and firm, grey. Smell and taste strongly farinaceous.
Spores 8.5-11(-12.5) x 7-10 um, Q = 1.0-1.3, almost isodiametrical, rather
obtusely angled. Basidia 4-spored. Lamellae edge fertile. Cystidia absent.
Pileipellis cutis of cylindrical hyphae, 4-13 um broad, pileitrama made of
inflated elements up to 22 um broad. Pigment yellow-brown intracellular.
Clamps abundant in all tissues. |
Hasitat: In dry heath land with various Mediterranean spiny scrubs
(Phrygana), on soil among Sarcopoterium spinosum scrubs.
COLLECTION EXAMINED: Greece, Isl. Amorgos, Kiklades (C. Aegean), Theologos
Monastery, 4 Dec. 2005, E. Polemis 05-M199.
Entoloma griseopruinatum is a very distinct species in stirps Lividoalbum in
subgenus Entoloma with relatively stout, firm basidiocarps. The dark colours,
non-hygrophanous, and pruinose pileus are distinctive (Noordeloos 2004). So
far it was known only from the type locality in France, Haute Savoie, where it
was found growing in a mountainous Fagus wood, a habitat very different from
the Greek location, which represents the second known locality of this rare
species. The type collection showed a distinct yellow tinge at the base of the
stipe, which has not been observed in the Greek material. Recently, it has been
recorded from two localities in Spain (Girona), in montane mixed broadleaved
forests of Corylus avellana, Betula pendula, Fraxinus excelsior, Salix caprea and
Sorbus aria (Vila & Caballero 2007). Entoloma griseorugulosum is somewhat
similar, differing, however, by a more fibrillose-rugulose pileal surface, and the
presence of incrusting pigment in the pileipellis (see below).
Entoloma griseorugulosum Noordel. & Fern. Sas. Fungi Europaei, 5a: 872 (2004) *
DESCRIPTION—Pileus 30-40 mm, convex then applanate with low umbo, margin
involute then straight, slightly lobed, not hygrophanous, not translucently
striate, dark grey-black to drab-sepia, becoming at centre smoke to mouse-
grey, silky, innately fibrous. Lamellae moderately distant, adnate, moderately
New Entoloma species from Greece ... 307
Figs. 3-7. Spores.
3. E. nigroviolaceum. 4. E. griseopruinatum. 5. E. griseorugulosum.
6. E. juncinum. 7. E. sordidulum. Bar = 10 um.
thick, smoke-grey to milky-coffee, with concolourous edge. Stipe up to 50 x
10-20 mm, ventricose, with somewhat rooting base, whitish-greyish, coarsely
fibrillose lengthwise, pruinose at apex, at base white tomentose, compact and
firm, with white rhizomorphs. Context brittle, grey, smell and taste distinctly
farinaceous.
Spores (9-)10-14(-17) x (8-)9.5-13.5(-15) um, Q=1-1.2, subisodiametrical,
with 6-8 obtuse to pronounced angles, moderately thick walled. Basidia 55-74
x 13-17 um, (2-)4-spored. Lamellae trama made up of cylindrical hyphae,
mostly longer than 100 um, with pustulate wall-surface. Lamellae edge fertile.
Cystidia absent. Pileipellis a cutis of long cylindrical, 3-8 um wide hyphae.
Pileitrama densely packed, made up of short inflated elements, in chains.
Pigment intracellular, and additionally incrusting in the narrow hyphae of
pileipellis and subpellis. Clamps abundant in all tissues.
308 ... Noordeloos & Polemis
HapsitTat: In calcareous Quercus ilex stand, on rich humus mixed with excessive
amounts of goat dung under Q. ilex.
COLLECTION EXAMINED: Greece, Isl. Naxos, Kiklades (C. Aegean), Kinidaros, 12 Dec.
2004, E. Polemis 04-N207.
Entoloma griseorugulosum was described from a thermophilic Quercus wood
in the Basque country, northern part of the Iberian peninsula, and is very
distinctive on account of its very dark basidiocarps, non-hygrophanous, strong
farinaceous smell, fibrous pileus and fibrous stipe surface, and two types of
pigment in the pileipellis.
Entoloma juncinum (Kihner & Romagn.) Noordel., Persoonia 10: 255 (1979)
DEscrIPTION—Pileus 20-40 mm, convex, soon plano-convex, obtuse, margin
slightly involute at first, but soon straight, slightly undulating, hygrophanous,
when wet dark brown at disc and striae, grey to milky-coffee towards the
margin, striate from the half radius up to disc, fading to yellowish-brown,
surface smooth, silky. Lamellae emarginated to almost free, beige then greyish-
pink. Stipe 60-70 x 2-5(-7) mm, cylindrical, compressed, broadened towards |
the base, concolorous with pileus, smooth and polished, apex pruinose.
Spores 7.5-10.5(-11) x 7-9(-10) um, Q = 1-1.2, almost isodiametrical, 5-7
angled. Basidia 32-42 x 10-13 um, 4-spored. Lamellae edge fertile. Cystidia
absent. Pileipellis cutis of cylindrical hyphae, to 12 um broad, with incrusting
yellow-brown pigment. Pileitrama made of inflated cells to 25 um broad, with
incrusted walls. Clamps present in hymenium, not seen elsewhere.
Hasirat: In a moist thicket by a stream, on humus-rich soil under Quercus
pubescens, Platanus orientalis and Alnus glutinosa (A361). Also in mountain
acidic heath land with Erica arborea, Genista acanthoclada, Crataegus monogyna,
and Pteridium aquilinum, on wet humus-rich, mossy soil (A413).
COLLECTIONS EXAMINED. Greece, Isl. Andros, Kiklades (C. Aegean), Evrousses, 22 Sept.
2002, E. Polemis 02-A361. Greece, Isl. Andros, Kiklades (C. Aegean), Vourkoti, 8 Oct.
2002, E. Polemis 02-A413.
Entoloma juncinum is a species with a very wide distribution, preferring
damp places in mixed forest on rather damp soil, but frequently also found in
grassland, where it can be confused with slender forms of Entoloma sericeum
Quél. (Noordeloos 2004).
Entoloma sordidulum (Kihner & Romagn.) P.D. Orton,
Trans. Br. Mycol. Soc. 43: 175 (1960)
DeEscripTioN—Pileus 2.5-4 cm, almost conical, with undulating margin,
slightly hygrophanous, when wet dark grey-brown, hazel at centre, olivaceous-
buff at margin, indistinctly translucently striate at extreme margin only, drying
New Entoloma species from Greece ... 309
to buff, surface smooth, innately fibrillose. Lamellae emarginate, greyish, then
vinaceous-buff to clay-pink, with smooth concolorous edge. Stipe 3-5 x 0.5-0.6
mm, cylindrical, greyish-brown smooth, silky-polished to indistinctly striate.
Context greyish-white, firm. Smell and taste rancid.
Spores 7—11(-12) x (7-)7.5-8.5(-9) um, Q=1.0-1.3, almost isodiametrical,
rather pronouncedly 5—6-angled in side-view. Basidia 4-spored. Lamellae edge
fertile. Cystidia absent. Pileipellis cutis of cylindrical hyphae, 3-10 um broad,
with minute to coarse incrusting and additionally some intracellular yellow-
brown pigment. Clamps rare in pileipellis, frequent in hymenium.
Hasirat: In thicket, on humus-rich soil under Quercus pubescens.
COLLECTION EXAMINED: Greece, Isl. Andros, Kiklades (C. Aegean), Falika, 10 Oct. 2002,
E. Polemis 02-A430.
Entoloma sordidulum is a widespread and common species, particularly in
deciduous and mixed forest on rich soil in North-western Europe (Noordeloos
2004). However, records from the Mediterranean are scarce. It is distinctive
with its relatively small and slender basidiocarps, dull brown colour, and
predominantly incrusting pigment in the pileipellis. It belongs to a group of
rather similar species in section Rhodopolia, which may be all mycorrhizal, and
difficult to distinguish.
Entoloma lividoalbum (Kihner & Romagn.) Kubi¢ka, Ceska Mykol. 29: 27 (1975)
DESCRIPTION—Pileus 90-110 mm, convex, then plano-convex with a blunt
but distinct umbo, finally applanate to somewhat depressed, margin at first
inflexed then straight and irregularly undulating, hygrophanous, indistinctly
translucently striate at the extreme margin, dark brown fading to light greyish-
beige, smooth, glabrous, with silky appearance. Lamellae moderately crowded,
emarginate, ventricose, white then clay-pink, with concolorous edge. Stipe
70-80(-90) x 10-20(-30) mm, cylindrical, attenuates towards the base,
whitish, fibrillose-striate lengthwise. Context thick, firm, white. Smell and taste
farinaceous.
Spores 9-11(-13) x 8-9(-11) um, Q=1.1-1.3, 5-7 angled, subisodiametrical.
Basidia 37-45 x 11-14 um, (2)4-spored. Lamella edge fertile. Cystidia absent.
Pileipellis an (ixo)cutis of narrow cylindrical hyphae. Pigment brown,
intracellular. Clamps abundant in all tissues.
Hasitat: In a thicket, on humus-rich soil under Quercus pubescens.
COLLECTION EXAMINED: Greece, Isl. Andros, Kiklades (C. Aegean), Agios Spyridon, 20
Dec. 1997, E. Polemis 97-A178.
Entoloma lividoalbum is a widespread species in Europe, and prefers Quercus
forest on rich soil. It is distinctive because of its relatively stout, tricholomatoid
habit, smooth, medium brown, hygrophanous pileus and white stipe.
310 ... Noordeloos & Polemis
Entoloma mougeotii (Fr.) Hesler, Beih. Nova Hedwigia 23: 158 (1967)
var. mougeotii
DESCRIPTION—Pileus 15-40 mm, convex with involute margin, slightly
umbilicate, expanding to applanate, with straight margin, not hygrophanous,
not translucently striate, initially dark violaceous-black, then dark grey,
violaceous-brown, uniformly coloured, or more often darker at the centre,
entirely tomentose to minutely squamulose, somewhat rimose-fissurate
when old. Lamellae adnate, or with a decurrent tooth, white then pink, with
concolorous, fimbriate margin. Stipe 20-60 x 1-4 mm, cylindrical to slightly
compressed, with longitudinally groove, steel-grey, at first longitudinally
fibrillose, then smooth shiny, white tomentose at base. Context thin, fragile at
stipe, easily splitting lengthwise. Smell indistinct and taste not recorded.
Spores (8.5-)9-11.5(-12) x 6-8 um, Q = 1.2-1.6, heterodiametrical, with
5-8 pronounced angles. Basidia 27-48 x 10-14 um, 4-spored. Lamella edge
sterile. Cheilocystidia 16-60(-80) x 7-17 um, cylindrical to slenderly clavate.
Pileipellis a trichoderm, made up of inflated cylindrical to clavate, or utriform
terminal elements, 6-25 um wide. Pigment violaceous-brown, intracellular.
Clamps absent from all tissues. |
Hasitat: Along a river's bank, on humus-rich soil under Platanus orientalis
and Alnus glutinosa (A446). On grassy soil, in field with Olea europaea and
Cupressus sempervirens.
COLLECTIONS EXAMINED: Greece, Isl. Andros, Kiklades (C. Aegean), Achlas River, 11
Oct. 2002, E. Polemis 02-A446. Greece, Isl. Andros, Kiklades (C. Aegean), Ypsilou, 28
Nov. 2002, E. Polemis 02-A618.
Entoloma mougeotit is distinctive because of its delicate violaceous-brown colour
and sterile lamella edge with large cheilocystidia. It has a wide distribution all
over Europe, but only in very special grassland habitats, for which it is a good
indicator species (Vesterholt 2002).
Entoloma cruentatum (Quél.) Noordel., Persoonia 12: 201. 1984.
DeEscripTION—Pileus 20-50 mm, hemispherical, strongly umbilicate,
with straight margin, expanding to convex, depressed, not hygrophanous,
translucently striate to the half of radius, more evidently as the pileus expands,
dark bluish to greyish black, at first uniformly coloured, then only at centre and
striae, with the rest of the surface greyish, smooth and silky, somewhat rugulose
and then adpressedly fibrillose at centre. Lamellae moderately distant, adnate,
greyish-blue, with concolorous margin. Stipe 30-50 x 2-3 mm, cylindrical to
slightly enlarged at base, concolorous with pileus, or somewhat paler, smooth,
polished, white tomentose at base.
Spores 9-12 x 7-9 um, Q = 1.3-1.5, heterodiametrical, with 6-8 pronounced
angles. Basidia 30-45 x 10-12 um, 4-spored. Lamella edge fertile. Cheilocystidia
New Entoloma species from Greece ... 311
absent. Pileipellis a trichoderm transiting to cutis towards the margin, made up
of inflated cylindrical to clavate, or utriform terminal elements, 7-20(-30) um
wide. Pigment brown, intracellular. Clamps absent from all tissues.
Habitat: In an open meadow with Olea europaea trees, on grassy soil.
COLLECTION EXAMINED: Greece, Isl. Andros, Kiklades (C. Aegean), Ypsilou, 15 Dec.
2007, E. Polemis 07-A906.
Entoloma cruentatum is very close to E. chalybeum, from which it mainly differs
by the fertile lamella edge without cheilocystidia. Usually a distinct orange
coloured mycelium has been observed at the base of the stipe, but this seems
to be absent in the Greek material. Entoloma cruentatum is a very rare species,
recorded so far from North-western Europe where it is a good indicator species
for old, poorly manured grasslands. It figures on the Red Data lists of Denmark,
Norway and Schleswig-Holstein (Vesterholt 2002). It has not been recorded
from the Mediterranean so far. A strongly umbilicate pileus, such as in our
specimens, has not been recorded for this species in pertinent descriptions.
Acknowledgments
The Moraitis Legacy and the Municipality of Korthi (Andros) are greatly acknowledged
for granting E. Polemis’ Ph. D. studies during 2003-2006. Access to the facilities
of the N.H.N (The Netherlands) for E. Polemis was achieved thanks to the financial
support from the SYNTHESYS Project http://www.synthesys.info/ which is financed
by European Community Research Infrastructure Action under the FP6 “Structuring
the European Research Area” Programme. Mrs Anita Walsmit-Sachs prepared the line
drawings for print, for which we are very grateful.
Literature cited
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4. Macrofungi from Abies cephalonica forests and from other intermixed tree species (Oxya Mt.,
central Greece). Mycotaxon 104: 39-42.
Gates G, Noordeloos ME. 2007. Studies in the genus Entoloma of Tasmania I. Persoonia 19:
157-226.
Manimohan P, Noordeloos ME, Dhanya AM. 2006. Studies on the genus Entoloma (Basidiomycetes,
Agaricales) in Kerala State, India. Persoonia 19: 45-93.
Noordeloos ME. 1980. Entoloma subgenus Nolanea in the Netherlands and adjacent regions with a
reconnaissance of list remaining taxa in Europe. Persoonia 10: 427-534.
Noordeloos ME. 1981. Introduction to the taxonomy of the genus Entoloma sensu lato (Agaricales).
Persoonia 11: 121-151.
Noordeloos ME. 1987. Entoloma (Agaricales) in Europe. Beihefte Nova Hedwigia 91. J. Cramer,
Vaduz.
Noordeloos ME. 1992. Entoloma s.|. Fungi Europei, vol. 5. Ed. Biella, Saronno.
Noordeloos ME. 2004. Entoloma s.l. Supplemento. Fungi Europei vol. 5a. Ed. Candusso, Alassio,
Italy.
312 ... Noordeloos & Polemis
Noordeloos ME, Hausknecht A. 2007. The genus Entoloma (Basidiomycetes, Agaricales) of the
Mascarenes and Seychelles, Fungal Diversity 27: 111-144.
Petrak F. 1943a.-Fungi. In KH Rechinger (ed.), Flora Aegaea, pp. 10-15.
Petrak F, 1943b. Fungi. In KH Rechinger (ed.), Neue beitrage zur flora von Kreta. Akad. Wissen.
Wien 105: 9-26
Plank S. 1980. Contribution to the study of wood-destroying fungi in Greece. Ann. Inst. Phytopathol.
Benaki 12: 247-256 [in Greek].
Polemis E, Noordeloos ME. 2007. Two new Gymnopus species from the Island of Andros (Kiklades,
C. Aegean, Greece). Mycotaxon 102: 171-178.
Polemis E, Zervakis GI, Dimou DM. 2002. New and interesting findings of macrofungi from
the islands of Andros and Naxos (Cyclades, Greece). Proceedings of the 7th International
Mycological Congress (11-17 August. 2002), Oslo, p. 169 (abstract).
Polemis E, Dimou DM, Tzanoudakis D, Zervakis GI. 2007. Rare ectomycorrhizal mushrooms
associated with Quercus coccifera from the Kiklades islands (Central Aegean, Greece). In the
proceedings of the XV Congress of European Mycologists (16-21 Sept. 2007), St Petersburg,
p. 140 (abstract).
Vesterholt J. 2002. Contribution to the knowledge of species of Entoloma subgenus Leptonia. Fungi
non delineati vol. 21. Editione Candusso, Alassio, Italy.
Vila J, Caballero F. 2007. Entoloma nuevos o interessantes de la Peninsula Iberica. Fungi non
delineati vol. 38. Editione Candusso Alassio, Italy. .
Zervakis G, Dimou DM, Balis C. 1998. A check-list of the Greek macrofungi including hosts and
biogeographic distribution: I. Basidiomycotina. Mycotaxon 66: 273-336.
Zervakis GI, Dimou D, Polemis E. 2004. Fungal diversity and conservation in the Mediterranean
area: recent advances in the inventory of the Greek macromycetes. Mycologia Balcanica 1:
31-34.
Zervakis G, Lizon P, Dimou D, Polemis E. 1999. Annotated check-list of the Greek macrofungi. II.
Ascomycotina. Mycotaxon 52: 487-506.
MY COTA XON
Volume 105, pp. 313-316 July-September 2008
A new species of Dactylellina producing adhesive knobs
and non-constricting rings to capture nematodes
HONGYAN Su’, Yu’E HAo?, XIAOYAN YANG?,
ZEFEN YU, JINGSHI DENG? & MINGHE Mo}
suhongyan16@yahoo.com.cn
* Department of Biology and Chemistry, Dali College
Dali 67000, P. R. China
* Department of Public Health of University of South China
Hengyang 421001, P. R. China
° Laboratory for Conservation and Utilization of Bioresources, Yunnan University
Kunming 650091, PR. China
Abstract — A nematode-trapping fungus, Dactylellina daliensis, is illustrated and
described as a new species. This fungus is primarily characterized by producing
elongate fusoid conidia with 1-13 (mostly 6-8) septa on the denticles of conidiophore
and capturing nematodes by means of stalked adhesive knobs and non-constricting
rings.
Key words — Antagonist of parasitic nematodes, predacious fungi, biological control,
trapping devices
Introduction
Nematode-trapping fungi are a group of hyphomycetes capable of capturing,
killing and digesting nematodes by special modified hyphal structures called
trapping devices. As antagonists of parasitic nematodes, these fungi have
been the subject of research over several decades. These fungi are abundant
in many habitats examined so far, with only a few species showing restricted
geographical distribution (Duddington 1951a, b; Gray 1987). During a recent
survey of the nematode-trapping hyphomycetes in rhizosphere soil of pine in
Yongping County, Dali Prefecture, Yunnan Province, a nematode-trapping
fungus with adhesive knobs and non-constricting rings was isolated and
identified. Comparing its morphological characters to the known species, and
following the concept of predacious hyphomycete genera as emended by Li et
al. (2005), we confirm that this species is a new one and name it as Dactylellina
daliensis.
314°4Su.& al.
Materials and methods
While surveying the predacious fungi, soil samples from pine rhizosphere
in Yongping County, Dali Prefecture, Yunnan Province were collected in
September 2007. Subsamples of 2-5 g were spread on Corn Meal Agar (CMA)
plates. After incubation at room temperature (about 20-28°C) for 10 days,
conidia of nematode-trapping fungi were transferred into fresh CMA plates
by sterile toothpicks to obtain pure cultures. For identification, cultures
obtained were inoculated on CMA and incubated at 28°C for 14-30 days and
the taxonomic characters were measured and determined. Conidial size and
septate were calculated by measuring more than 100 conidia. Microscopic
photographs were taken using an Olympus BX51 microscope. To induce trap
formation, a 2 cm? piece of agar in the center of the plate was removed from a
7-day old culture to create an open space. About 200 nematodes (Panagrellus
redivivus) were added into the free space after the mycelia emerged from the
cut margin.
Taxonomic description
Dactylellina daliensis Hong Yan Su, sp. nov. PLATE 1
MycoBank MB 511994
Coloniae albae in CMA, ad 6 cm diam. post 7 dies 25 °C. Mycelii sparsa, hyphae hyalinae,
septata, ramosa, 2.2-3.7 um lata. Conidiophora erecta, hyalina, ramosa, 2-4 septata, 21-
101 ym longa, 1.8-2.5 um crassa ad basem, 1.5-1.7 um crassa ad apicem, lata conidla
greges in perspicuis dendriculis in capitata. Conidia hyalini, elongati fusiformis vel clavati,
1-13 septati, praecipue 6-8 septati, 28.7-51.5 (40.8) x 3.2-5.2 (4.2) um.
Ho.LotyPeE: YMFT 1.0213, permanent slide, Yongping County, Dali Prefecture, Yunnan
Province, P.R. China, HongYan Su, Sep. 2007.
ETYMOLOGY: species epithet refers to the collection site of the species.
Colonies white on CMA, growing rapidly, obtaining a diameter of 6 cm at 25°C
within 7 days. Mycelium sparse, hyphae hyaline, septate, branched, 2.2-3.7 um
wide. Conidiophores erect, colorless, more or less branched, 2-4 septate, 21-101
uum high, 1.8-2.5 um wide at the base, gradually tapering upward to a width
of 1.5-1.7 um at the tip where forming 1-7 apical conidia. Conidia hyaline,
elongated fusoid-shaped or clavate, straight or somewhat curved, round at the
distal end, truncate at the base, 1-13 (mainly 6-8) septate, 28.7-51.5 (40.8) x
3.2-5.2 (4.2) um in length and width. Trapping devices, stalked adhesive knobs
and non-constricting rings, forming on water agar medium when challenging
with living nematodes. Chlamydospores not observed in culture.
Discussion
In the known species of nematode-trapping fungi, Dactylellina daliensis only
resembles only three other species in the genus: D. haptospora (Drechsler
. OED
Dactylellina daliensis sp. nov. (China)
Plate 1. Dactylellina daliensis. A-B. Conidiophores and conidia on CMA _ medium.
C-G. ‘Conidiophores. H-P. Conidia. Q. Stalked knobs on conidia and mycelia.
R. Non-constricting rings. Scale bars: A=50 um, B=25um, C, E=12.5 pm, D, F-R=5 pm.
316... Su & al.
1940), D. leptospora (Drechsler 1937) and D. varietas (Li et al. 2006). Both
D. daliensis and D. leptospora produce elongated fusoid conidia and capture
nematodes by the same kinds of trapping devices (stalked adhesive knobs and
non-constricting rings). However, unlike D. daliensis, which forms terminal
cluster of conidia, D. leptospora mainly bears a single terminal conidium,
occasionally producing an additional conidium on a short branch attached to
the main axis near the conidiophore apex. Furthermore, the conidial length of
D. daliensis (28.7-51.5 um) is shorter than that of D. leptospora (40-105 um).
D. daliensis and D. varietas have similar conidial shape, arrangement of conidia
on conidiophores, and trapping device types, but they differ in the size of
conidia, 28.7-51.5 (40.8) x 3.2-5.2 (4.2) um for D. daliensis and 25-61.5 (46.5)
x 6.5-10 (9) um for D. varietas. Additionally, unlike D. varietas, conidia of
D. daliensis can form stalked adhesive knobs on their terminal. Both D. daliensis
and D. haptospora can bear terminal conidial clusters and form stalked adhesive
knobs as trapping devices. However, D. daliensis differs from D. haptospora by
producing an additional type of trapping device, non-constricting rings.
Acknowledgements
This work was supported jointly by “National Basic Research Program of China’
(2007CB116310), 2005C0076M, 06Y128B and 2005Q007B. We are grateful to Dr.
ShiDong Li and Dr. XueFeng Liu for revising the manuscript and valuable comments.
We are also indebted to Prof. Shugang Lu for helping with the Latin.
Literature cited
Drechsler C. 1937. Some hyphomycetes that prey on free-living terricolous nematodes. Mycologia
29: 447-552.
Drechsler C. 1940. Three new hyphomycetes preying on free-living terricolous nematodes.
Mycologia 32: 448-470.
Duddington CL. 1951a. The ecology of predacious fungi. I. Preliminary survey. Trans. Brit. Mycol.
Soc. 34: 322-331.
Duddington CL. 1951b. Further records of British predacious fungi. I. Trans. Brit. Mycol. Soc. 34:
194-209.
Gray NE. 1987. Nematophagous fungi with particular reference to their ecology. Biol. Rev. 62:
245-304.
Li Y, Hyde KD, Jeewon R, Cai L, Vijaykrishna D, Zhang KQ. 2005. Phylogenetics and evolution of
nematode-trapping fungi estimated from nuclear & protein coding genes. Mycologia 97(5):
1034-1046.
Li Y, Jeewon R, Hyde KD, Mo MH, Zhang KQ. 2006. Two new species of nematode-trapping fungi:
relationships inferred from morphology, rDNA and protein gene sequence analyses. Mycol.
Res. 110: 790-800.
MYCOTAXON
Volume 105, pp. 317-323 July-September 2008
Studies in Amanita subsection Vittadiniae |—
a new species from Colombian savanna
RODHAM E. TULLOSS!
< ret@eticomm.net >
P. O. Box 57, Roosevelt, New Jersey 08555-0057, USA
A. E. FRANCO-MOLANO
< afranco@quimbaya.udea.edu.co >
Laboratorio de Taxonomia y Ecologia de Hongos
Instituto de Biologia, Universidad de Antioquia
A.A. 1226, Medellin, Colombia
This paper is dedicated to DR. CORNELIS BAS on the occasion
of his eightieth birthday.
Abstract—Amanita savannae is described as new from the Departamento de Meta,
Colombia.
Key words—Amanita subsect. Limbatulae, Amanita subsect. Solitariae, South America,
taxonomy
Introduction
Amanita savannae is a member of the growing group of apparently nonmyc-
orrhizal, or at least not obligately mycorrhizal, species in Amanita subsect.
Vittadiniae Bas. The paper of Wartchow et al. (2007) can be considered a pre-
vious work in the present series. Previous work on Amanita of Colombia
includes (Tulloss et al. 1992). Illustrations and brief descriptions of Colom-
bian species of Amanita as well as other taxa can be found in (Franco-Molano
et al. 2000).
Materials and methods
Methods used are those of Bas 1969 and Tulloss et al. (1992) as modified by
Tulloss (1993, 1994, 2000, 2008). Color codes of the form 4A5 follow
Kornerup and Wanscher (1978). Herbarium name abbreviations are confor-
|. Research Associate (hons.), the New York Botanical Garden (NY).
318...Tulloss & Franco-Molano
Fig. 1. Amanita savannae (holotype). Habit (<1.7), reconstructed from exsiccatum. Pencil
sketch by Richard Rauh (NY).
mant with (Holmgren et al. 1990), with the exception of RET (Tulloss’ her-
barium). Abbreviations in author citations follow Kirk and Ansell (1992,
2008).
Taxonomic part
Amanita savannae Tulloss & Franco-Mol. sp. nov. Fig. 1
MYCOBANK # 511862
Pileus 12-30 mm latus, albus vel griseobrunneus, convexus vel planus vel
concavus, squamellulis tenuibus et brunneis insidens disci. Lamellae albae.
Stipes 13-30 x 4—7 mm, pallidus, annulo albo, membranaceo, medio vel infe-
riori, bulbo subnapiformi. Fibulae praesentes. Sporae subglobosae vel late
ellipsoideae, amyloideae, interdum globosae, interdum elongatae, (6.5—) 7.2—
10.8 (-13.5) x (6.0—) 6.2-8.5 (—9.5) um. Habitat in savanna humida. In sub-
sectione Vittadiniis stirpis Vittadinii amplitudinae basidiocarpi, praesentia et
forma bulbi basilaris stipitis, praesentia particula dextrinoidea in basidiis, et
amplitudine et forma sporarum distinguenda. Holotypus: Colombia, Dpto.
del Meta, Mpio. Puerto Gaitan, Puente Arimena, sobre la carretera a Carima-
gua, 18.v.1992 A. E. Franco-Molano 850 (NY).
ETYMOLOGY: “‘of the savanna.”
PILEUS: 12-30 mm wide, white to grayish brown (6B2), convex to planar to
concave; context white, unchanging when cut or bruised, 2+ mm thick at stipe;
margin nonstriate, appendiculate, decurved to inflexed at first, eventually flar-
Amanita from Colombia...319
ing upward, entire; wniversal veil as fine squamules, densest over disk, brown
(6ES).
LAMELLAE: free, subclose, white, unchanging when cut or bruised, ventricose,
not marginate, 2* mm broad; /amellulae attenuate, of at least two lengths.
STIPE: 13-30 (including bulb) x 4—7 mm, pallid, narrowing upward or down-
ward or subcylindric, smooth and glabrous toward the apex, covered with
rather fine and pallid fibrillose scales below annulus; bu/b subnapiform; con-
text white, unchanging when cut or bruised, solid; partial veil white, membra-
nous, median to inferior, sometimes left in part on pileus margin or covering
the outer portion of lamellae; universal veil distributed on the lower stipe as
scales (see above) and in broken ring of pallid tissue around apex of bulb.
Odor mild. Taste not recorded.
MACROCHEMICAL TESTS: Spot test for tyrosinase (paracresol) - positive
throughout basidiocarp after 30 min. Spot test for laccase (syringaldazine) -
negative throughout basidiocarp. Test voucher: Franco-Molano 850.
PILEIPELLIS: lacking; pileus context intergrading with universal veil through
region with elements more closely packed than lower in pileus context.
PILEUS CONTEXT: filamentous, undifferentiated hyphae 1.2—9.8 um wide,
intergrading with narrow inflated cells, occasionally having intercalary
inflated cells similar to the acrophysalides of this tissue, branching, walls thin
or slightly thickened as in the acrophysalides; acrophysalides subfusiform to
narrowly fusiform to clavate to fusiform-rostrate to narrowly clavate to elon-
gate-ellipsoid, up to 169 <x 26 um, dominating, sometimes branching, with
walls thin or thickened (up to 0.8 um thick), often with subradial orientation;
vascular hyphae 3.2—8.0 um wide, branching, common; clamps observed.
LAMELLA TRAMA: bilateral, with central stratum (W,, = 25-35 um) appearing
dark and quite distinct at low magnification, with angle of divergence from
very shallow to nearly perpendicular to the central stratum; filamentous,
undifferentiated hyphae 1.8—6.0 um wide, branching; terminal, inflated cells
not observed; vascular hyphae not observed. SUBHYMENIUM: a branching
structure of short uninflated hyphal segments, with wg-near = 15 um and we-
far = 45 um, rather small partially inflated to inflated (e.g., 10 < 8.5 um) inter-
calary cells, and irregular (e.g., branched) elements, densely interwoven, with
elements approaching bases of basidia at 45—90° angle, with most basidia aris-
ing from uninflated hyphal segments, but some (especially in regions that are
generally more inflated) arising from small inflated cells. BASIDIA: 32-64 x
10.0-14.0 um, 4-sterigmate, containing dextrinoid granules; clamps rather
plentiful. UNIVERSAL VEIL: On pileus: extensively gelatinized, more yellow-
brown than adjacent pileus context, with elements having periclinal orienta-
320...Tulloss & Franco-Molano
tion, with most extensively gelatinized regions red-brown and amorphous; fil-
amentous, undifferentiated hyphae 2.5—6.2 um wide, branching; inflated cells
in chains, dominant, subfusiform to narrowly clavate, up to 53 x 21 um, all
badly damaged in material reviewed; vascular hyphae not distinguished due to
extensive gelatinization. On stipe base: (from broken ring at bulb apex)
slightly sordid in mass; filamentous, undifferentiated hyphae 3.0-8.5 um
wide, branching, plentiful; inflated cells in easily dissociated chains, domi-
nant, elongate to cylindric to clavate to ellipsoid, up to 99 x 29 um, with walls
thin to slightly thickened to 0.8 um thick; vascular hyphae not observed;
clamps present. STIPE CONTEXT: longitudinally acrophysalidic; filamentous,
undifferentiated hyphae 2.2—8.8 um wide, dominating, branching; acrophysal-
ides subfusiform to clavate, rather slender, up to 158 x 45 um, with walls thin
or slightly thickened; vascular hyphae 5.0—18.5 um wide, branching; clamps
present. PARTIAL VEIL: upper surface partially gelatinized; filamentous,
undifferentiated hyphae 1.0—6.5 um wide, branching, dominating, criss-cross
interwoven; inflated cells terminal, unevenly distributed, subclavate to sub-
fusiform, up to 114 x 36 um, with slightly thickened walls; vascular hyphae
not observed; clamps present.
BASIDIOSPORES: [140/7/1] (6.5—) 7.2—10.8 (—13.5) x (6.0—) 6.2—8.5 (—9.5) um,
(L = 7.6—9.2 (—10.2) um; L’ = 8.6 um; W = 6.6—7.6 (—8.2) um; W’ = 7.2 um; Q
= (1.02—) 1.08—-1.36 (-1.73); Q = 1.14-1.22 (—1.27); Q’ = 1.19), hyaline,
smooth, thin-walled, amyloid to somewhat palely amyloid, subglobose to
broadly ellipsoid to ellipsoid, occasionally globose, occasionally elongate,
often not adaxially flattened; apiculus sublateral, proportionately small to very
small, cylindric; contents multiguttulate; color in deposit unknown.
ECOLOGY : Subgregarious to gregarious. At 120 m elev. In wet savanna,
apparently without nearby woody plants, recently flooded by heavy rains.
MATERIAL EXAMINED: COLOMBIA: EL META—Mpio. Puerto Gaitan - Puente
Arimena, sobre la carretera a Carimagua, 18.v.1992 A. E. Franco-M. 850 (holotype,
NY; isotype, COL; isotype, RET).
NOTES: This taxon is easily keyed to Amanita stirps Vittadinii using the keys
of Bas (1969). The species most similar to A. savannae that is treated by Bas
(1969) is A. codinae (Maire) Bertault (1955), described from the Mediterra-
nean region. However, A. codinae has a rather thickset habit (pileus up to 130
mm wide) with a roughly cylindric stipe (up to 80 mm long) that lacks both a
bulb and a point at the bottom; and it has much larger spores than does A.
savannae with Q between 1.45 and 1.60.
Amanita from Colombia...321
Fig. 1. Amanita savannae (holotype). a. Elements of gelatinized universal veil on pileus. b.
Elements of universal veil from stipe base. c. Exsiccatum, stipe base. d. Elements of hyme-
nium and subhymenium. e. Elements of pileus context subjacent to universal veil. f. Elements
of partial veil. For stipe base, scale bar = | cm, for all microscopic drawings, = 20 1m.
322...Tulloss & Franco-Molano
Described from the State of Parana, southern Brazil, Amanita grallipes Bas &
de Meijer (1993) differs in being. larger (pileus up to 90 mm wide; stipe up to
120 mm long) with downward tapering stipe lacking a bulb, lamellae becom-
ing golden yellow (4A6) in age, and smaller spores with higher Q value.
Among the remaining taxa of subsect. Vittadiniae described since the publica-
tion of (Bas 1969), the only other taxon with which A. savannae might be con-
fused is A. pleropus (Kalchbr. & MacOwan) D. A. Reid (1975) described from
South Africa (Reid & Eicker 1991, 1996). However, there are marked differ-
ences between the two species. Amanita pleropus has a universal veil with
elements having anticlinal orientation. As a result the universal veil remnants
on the pileus sometimes become “densely crowded, erect, hair-like spines or
pyramidal warts.” The African species also has larger spores (10.8—14.0 x
7.0—9.5 um) with Q in the range of 1.5*.
Dextrinoid (or possibly dextrinoid) contents of basidia or spores is a character
reported for two other species in Amanita sect. Lepidella—A. mutabilis
Beardslee (Bas 1969, Tulloss 1984), belonging to subsect. Limbatulae Bas
and A. westii (Murrill) Murrill (Bas 1969, Tulloss and Lewis 1994), belonging
to subsect. Solitariae Bas. This paper provides the first report of dextrinoid
granules in Amanita subsect. Vittadiniae. These granules apparently occur
rarely outside of Amanita subgen. Lepidella (E.-J. Gilbert) Beauseign. emend.
Corner & Bas; they have been noted by Tulloss (unpub. data) in A. umbrino-
lutea var. flaccida D. A. Reid (1987) of Amanita sect. Vaginatae (Fr.) Quél.
sensu Yang (1997) and others. Such granules were considered of taxonomic
importance in the case of A. mutabilis; however, whether or not they are
genetically determined is an open Issue.
Acknowledgments
We gratefully acknowledge the assistance and support of the following persons: Dr. Z.
L. Yang, Academia Sinica, Heilongtan, Kunming, Yunnan, China, and Dr. M. Kuo,
Charleston, Illinois, USA, reviewed this article during its preparation for publication.
Dr. S. R. Pennycook, Landcare Research, Auckland, New Zealand, provided a thor-
ough nomenclatural review and provided valued corrections and related correspon-
dence. Mr. Richard Rauh (NY) prepared our habit illustration. Mrs. Mary A. Tulloss,
Roosevelt, New Jersey, assisted in preparation of this paper for publication. Franco-
Molano’s field work in Colombia was made possible by a Burlingham Fellowship in
Mycology at The New York Botanical Garden.
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Amanita from Colombia...323
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Tulloss RE. 1984. Distribution and taxonomic notes on Amanita mutabilis. Mycologia 76: 555—
558.
Tulloss RE. 1993. Amanita pachysperma, Amanita subvirginiana, and Amanita virginiana (taxon-
omy and distribution) with notes on description of the lamella trama in Amanita. Mycotaxon 49:
449-475.
Tulloss RE. 1994. Type studies in Amanita section Vaginatae |: Some taxa described in this Cen-
tury (studies 1-23) with notes on description of spores and refractive hyphae in Amanita. Myco-
taxon 52: 305-396.
Tulloss RE. 2000. Note sulla metodologia per lo studio del genere Amanita (Agaricales). Boll.
Gruppo Micol. G. Bresadola 43(2): 41-58.
Tulloss RE, Lewis DP. 1994. Amanita westii—taxonomy and distribution. A rare species from
states bordering on the Gulf of Mexico. Mycotaxon 50: 131-138.
Tulloss RE, Ovrebo CL, Halling RE. 1992. Studies on Amanita (Amanitaceae) from Andean
Colombia. Mem. New York Bot. Gard. 66: 1-46.
Wartchow F, Tulloss RE, Cavalcanti MAQ. 2007. The discovery of Amanita lilloi in Brazil.
Mycotaxon 99: 167-174.
Yang ZL. 1997. Die Amanita-Arten von Siidwestchina. Biblioth. Mycol. 170: 1-1, 1-240.
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7
MYCOTAXON
Volume 105, pp. 325-330 July-September 2008
Zygophiala (hyphomycetes) —
a genus newly recorded from China
XIAORU ZHAI’, HUANYU Lr, RONG ZHANG! , GUANGYU SUN",
MING TANG’, JEAN C. BATZER? & MARK L. GLEASON?
sgy@nwsuaf.edu.cn
‘College of Plant Protection
Yangling, Shaanxi, 712100, China
* College of Forestry, Northwest AéF University
Yangling, Shaanxi, 712100, China
, ‘Department of Plant Pathology, Iowa State University
Ames, Iowa 50011, U.S.A
Abstract — We document the first report of the genus Zygophiala from China. The
fungus Zygophiala wisconsinensis was isolated from colonies on the cuticle of apple
fruit collected from 2 orchards in Shaanxi and Henan Provinces. Conidiophores were
erect, scattered, subcylindrical, irregularly flexuous, s-shaped, consisting of four parts:
a hyaline supporting cell that gives rise to a smooth, dark brown stipe, terminating in
a finely verruculose, medium brown apical cell that gives rise to two or one medium
brown, finely verruculose, doliiform to ellipsoidal, polyblastic conidiogenous cells,
with 1-2 prominent scars, apical and lateral, darkened, thickened. Conidia were
solitary, fusiform to obclavate, hyaline, smooth and thick-walled, transversely 1-septate,
prominently constricted at the septum; apex obtuse, base subtruncate, with a darkened,
thickened hilum.
Key words —flyspeck, internal transcribed spacer, rDNA, phylogenetic analysis
Introduction
Flyspeck is a complex of fungi that blemishes the cuticle of pome fruits
throughout moist temperate regions of the world. Colby (1920) reported that
flyspeck was caused by Leptothyrium pomi (Mont. & Fr.) Sacc. Von Arx (1959)
synonymized 14 species, including Leptothyrium pomi, to Schizothyrium pomi
(Mont. & Fr.) Arx., the currently accepted name. Martyn (1945) reported
Zygophiala jamaicensis E.W. Mason as the pathogen of leaf speckle of banana in
Jamaica. The link between S. pomi and its Zygophiala anamorph was reported
*Corresponding author.
326 ... Zhai & al.
by Durbin et al. (1953), who inoculated immature apple fruit with ascospores,
which produced both the teleomorph and anamorph states. Schizothyrium pomi
and its Zygophiala anamorph were reported to cause greasy blotch of carnation
and flyspeck of apple in the United States (Baker et al. 1977). Evidence that
S. pomi and Z. jamaicensis are the same species remains anecdotal, however.
Baines (1940), Baker et al. (1977) and Nasu & Kunoh (1987) reported that
Z. jamaicensis occurred on 120 species in 44 families of plants. Based on
surveys of apple orchards in the midwestern and eastern U.S., Batzer et al.
(2008) described four new species of the genus Zygophiala. They suggested that
S. pomi might not be the teleomorph of Z. jamaicensis, based on difference in
size of conidia between the anamorph of S. pomi and Z. jamaicensis.
Materials and methods
In September 2006, apples with flyspeck signs were collected from 2 orchards;
isolate BJ34 was from Baoji County, Shaanxi Province, and isolate LB15
was from Lingbao County, Henan Province. The thalli were transferred to a
potato dextrose agar (PDA) slant directly from the apple fruit surface, and.
then incubated at 25°C in darkness (Sun et al. 2003). The characteristics of
2- to 4-wk-old cultures on PDA and on OA were described and photographed
(Fig. 1).
Table 1 Sequences downloaded from GenBank
SEE SS ‘TELEOMORPH ANAMORPH REFERENCE
AY598854 Zygophiala cryptogama Batzer et al. 2008
EF164900 Z. cryptogama Batzer et al. 2008
EF164899 Z. cryptogama Batzer et al. 2008
AY598848 Schizothyrium pomi Batzer et al. 2008
EF164898 S. pomi Batzer et al. 2008
AY598850 S. pomi Batzer et al. 2008
AY598853 Z. wisconsinensis Batzer et al. 2008
AY598855 Z. wisconsinensis Batzer et al. 2008
AY598856 Z. tardicrescens Batzer et al. 2008
AF173313 Mycosphaerella molleriana — Crous et al. 1999
Zygophiala new to China... 327
POLYMERASE CHAIN REACTION AND SEQUENCING. —Template DNA was
extracted according to the method of Barnes et al. (2001), and primer pairs
used for amplification and sequencing of the ITS region were ITS1-F (Gardes
et al. 1993) and ITS4 (White et al. 1990). Amplification was completed with
the following cycling parameters: initial denaturation at 94°C for 3 min
followed by 35 cycles of denaturation at 94°C for 30 s, annealing at 52°C for
30 s, and extension at 72°C for 10 min. The PCR products were sequenced by
the Organism Technology Co., Ltd., Shanghai, China. Accession Number in
GenBank is EU329727 for isolate BJ34, EU825775 for isolate LB15.
SEQUENCE ALIGNMENT AND PHYLOGENETIC ANALYSIS. —The ITS nucleotide
sequences generated in this study were added to other sequences downloaded
from GenBank (Table 1), with high homology according to a BLAST search
(National Center for Biotechnology Information’s nucleotide blast program).
Preliminary alignments were performed using CLUSTAL-X. The alignments
were imported into BioEdit 5.0.9.1 and manually adjusted. Phylogenetic analysis
of aligned DNA sequences was performed with PAUP version 4.0b10 for 32-bit
Microsoft Windows (Swofford 2001). Heuristic searches were performed with
1000 random sequence additions. The clade stability was evaluated by 1000
bootstrap replications. Other measures for parsimony, including tree length,
consistency index, retention index and rescaled consistency index (CI, RI and
RC, respectively), were also calculated. Mycosphaerella molleriana was used as
the outgroup taxon.
Results
On the surface of apple fruit, the fungus produced dark, shiny, round to oval,
slightly protuberant sclerotium-like bodies, 344 (286-357) x 473 (357-607)
um.
Co.onigs on OA when 2- to 4-week old spreading with aerial mycelium and
smooth, regular margins; the middle of colonies olivaceous gray, outer zone pale
white; diameter of 2-week growth at 25°C 5 mm. CONIDIOPHORES arising from
superficial hyphae, 2-3 um wide, scattered, 1-septate, subcylindrical, irregularly
flexuous, consisting of a hyaline supporting cell that gives rise to a smooth,
dark brown stipe, 15-20 x 4-6 um, (from basal septum to below phialide);
terminating in a finely verruculose, medium brown apical cell, 3-5 x 4-6 um,
that gives rise to two or one medium brown, finely verruculose, doliiform to
ellipsoidal, polyblastic conidiogenous cells, 5—7 x 4-5 um, with 1-2 prominent
scars, apical and lateral, darkened, thickened, somewhat refractive, 2-3 um
wide. Conip1A solitary, fusiform to obclavate, hyaline, smooth and thick-
walled, granular, aseptate, 14-22 x 6-9 um, or transversely 1-septate (rarely
median), prominently constricted at the septum; apex obtuse, base subtruncate,
with a darkened, thickened hilum, 2-3 um wide.
328 ... Zhai & al.
_
.
a SS
ts
Fig. 1 Zygophiala wisconsinensis BJ34
A-C, conidia and conidiophores; D, colony on PDA; bars (A-C) = 3 um, bars (B) = 5 um
ITS sequences were generated from two isolates (BJ34, LB15) obtained from
the cuticle of apple fruit. Ten GenBank ITS sequences were aligned with the
two isolate sequences to generate a majority consensus tree (Fig. 2) with 90
length (CI = 0.9140, RI = 0.8788, RC = 0.8032). One clade with 77% bootstrap
support included several strains of S. pomi and Z. cryptogama Batzer & Crous;
the second clade had 79% bootstrap support. Isolates BJ34 and LB15 grouped
with Zygophiala wisconsinensis Batzer & Crous with 94% bootstrap support.
rar ne a
oe
Zygophiala new to China... 329
AY598854 Zygophiala cryptogama
89 EF 164900 Z cryptogama
77 EF164899 Z cryptogama
AY598848 Schizothyrium pomi
78 EF 164898 S. pomi
AY598850 S. pomi
AY598853 Z wisconsinensis
94 AY598855 Z wisconsinensis
LB15
79
BJ34
AY598856 Z ftardicrescens
AF 173313 Mycosphaerella moileriana
Fig. 2 The majority consensus tree (length = 93, CI = 0.9140, RI = 0.8788, RC = 0.8032) derived
from a heuristic search option in PAUP version 4.0b10 for 32-bit Microsoft Windows with 1000
randomizations of sequence input orders and 1000 bootstrap replications using the data set of
ITS1, 5.8S and ITS2. Bootstrap values > 50% are indicated above the tree branches.
Discussion
This is the first report of the genus Zygophiala and of Z. wisconsinensis on
apple in China. Zygophiala spp. can be distinguished by colony and conidial
morphology. Z. tardicrescens Batzer & Crous can be differentiated from
Z. wisconsinensis by diameter growth on MEA. The sclerotium-like bodies of
Z. wisconsinensis on pome fruit also tend to be ovoid, larger, and more sparsely
arranged than for other flyspeck species (Batzer et al. 2008). Our research
indicates that strains isolated from the apple cuticle grew more quickly on
PDA than on OA, with conidial morphology and size matching those of
Z. wisconsinensis. DNA sequence analyses of both isolates also demonstrate
their close relationship with Z. wisconsinensis. Based on these characters we
identified the isolates as Z. wisconsinensis.
330 ... Zhai & al.
Acknowledgments
This work was supported by National Natural Science Foundation of China (30670013
and Key Project No. 30630054) and Program for Changjiang Scholars and Innovative
Research Team in University (No. 200558 and IRT0748). The authors wish to thank
Professors Turner B. Sutton (North Carolina State University, Raleigh, NC, U.S.A.) and
Zhongyi Zhang (College of Plant Protection, Yunnan Agricultural University, Kunming,
Yunnan, China) for reviewing the manuscript.
Literature cited
Baines RC. 1940. Pathogenicity and hosts of the flyspeck fungus of apple. Phytopathology 30: 2.
Baker KF, Davis LH, Durbin RD, Snyder WC. 1977. Greasy blotch of carnation and flyspeck disease
of apple: diseases caused by Zygophiala jamaicensis. Phytopathology 67: 580-588.
Barnes I, Gaur A, Burgess 'T, Roux J, Wingfield BD, Wingfield MJ. 2001. Microsatellite markers
reflect intra-specific relationships between isolates of the vascular wilt pathogen, Ceratocystis
fimbriata. Molecular Plant Pathology 2: 319-325.
Batzer JC, Arias MMD, Harrington TC, Gleason ML, Groenewald JZ, Crous PW. 2008. Sooty blotch
and flyspeck fungi on apples. Four species of Zygophiala (Schizothyriaceae, Capnodiales) are
associated with the sooty blotch and flyspeck complex on apple. Mycologia 100(2): 232-244.
Colby AS. 1920. Sooty blotch of pomaceous fruits. Trans. Ill. Acad. Sci. 13: 139-179.
Durbin RD, Davis LH, Snyder WC, Baker KF. 1953. The imperfect stage of Mycothyriella rubi, cause
of flyspeck of apple. Phytopathology 43: 470-471.
Gardes M, Bruns TD. 1993. ITS primers with enhanced specificity for basidiomycetes — application
to the identification of mycorrhizae and rusts. Mol. Ecol. 2: 113-118.
Martyn EB. 1945. Note on banana leaf speckle in Jamaica and some associated fungi. Mycol. Pap.
13: 1-5.
Nasu H, Kunoh H. 1987. Distribution of Zygophiala jamaicensis in Okayama Prefecture, Japan.
Trans. Mycol. Soc. Jpn. 28: 209-213. .
Sun GY, Zhang R, Zhang Z, Zhang M. 2003. Isolation of sooty blotch and flyspeck fungi from apple
surface by picking up the thalli. Acta Phytopath. Sin. 33 (5): 479-480 [in Chinese].
Swofford DL. 2001. PAUP* Phylogenetic analysis using parsimony (*and other methods) version
4.0 beta version. Sinauer Associates, Sunderland, Massachusetts, U.S.A.
White TJ, Bruns TD, Lee SB, Taylor JW. 1990. Analysis of phylogenetic relationships by amplification
and direct sequencing of ribosomal RNA genes. In: Innis MA, Gelfand DH, Sninsky JJ, White TJ
(eds) PCR protocols: a guide to methods and applications. Academic, New York, pp 315-322.
MYCOTAXON
Volume 105, pp. 331-336 July-September 2008
Two new species and a new Chinese record of Exobasidium
(Exobasidiales) from China
ZHENYING LI’? & LIN Guo'*
lizheny@mails.gucas.ac.cn *guol@sun.im.ac.cn
'Key Laboratory of Systematic Mycology and Lichenology
Institute of Microbiology, Chinese Academy of Sciences
Beijing 100101, China
?Graduate University of Chinese Academy of Sciences
Beijing 100049, China
Abstract—Two new species, Exobasidium rhododendri-nivalis on Rhododendron nivale
and E. pyroloides on Gaultheria pyroloides, are reported. They were collected from
Yunnan and Sichuan Provinces. Exobasidium rhododendri-nivalis causes small galls
on leaves, stems and shoots, while E. pyroloides causes red leaf spots. Exobasidium
cylindrosporum on Rhododendron sp., collected from Jiangxi Province, is reported as
new to China.
Key words— Ustilaginomycetes, symptoms, taxonomy
Two new species of Exobasidium, collected from southwestern China, are
described and illustrated. The first new species was collected in 2007 from
Yunnan and Sichuan Provinces at altitudes of 4300 m and 4650 m. It is parasitic
on Rhododendron nivale (subfamily Rhododendroideae of Ericaceae), causing
small galls measuring 1-4 mm in diam. on leaves, stems and shoots. On leaves
there are 1-5 (or more) galls on the lower surface. Diseased leaves are convex
on the upper surface. The galls are red when fresh and become pale yellowish
brown to black when old. Basidiospores with short germ tubes were observed
in some microscopical slides of fresh material. The new species is described
as:
Exobasidium rhododendri-nivalis ZhenYing Li & L. Guo, sp. nov. Figs. 1, 4-7
MycoBank MB 511910
Hymenium album. Basidia clavata vel cylindrica, 7-18 x 4-8 um, hyalina, terminaliter
2-4 sterigmatibus 4.5-7 x 1-1.5 um praedita. Basidiosporae ellipsoideae, curvae, 10.2-13
x 2.5-3 um, hyalinae, laeves, primo continuae, dein 1-septatae; per hyphas germinantes.
*corresponding author
352. eb PecGuo
Fig. 1. Line drawings of basidia, sterigmata and basidiospores of
Exobasidium rhododendri-nivalis on Rhododendron nivale (HMAS 183431, holotype).
Hymenium white, forming a continuous layer over the surface of a gall.
Basidia clavate or cylindrical, 7-18 x 4-8 um, hyaline, with 2-4 sterigmata.
Sterigmata conical, 4.5-7 x 1-1.5 um. Basidiospores ellipsoidal, 10.2-13 x
2.5-3 um, hyaline, smooth, at first continuous, then 1-septate, and slightly
curved; germinating by germ tubes.
SPECIMENS EXAMINED—On Rhododendron nivale Hook.f. (Ericaceae), Yunnan: Degen,
Baimaxue Mountain, alt. 4300 m, 21 IX 2007, Z.Y. Li, L. Guo & S.H. He 697, HMAS
183431 (holotype); Sichuan: Xiangcheng, alt. 4650 m, 24 IX 2007, Z.Y. Li, L. Guo & S.H.
He 716, HMAS 183444 (paratype)
Exobasidium rhododendri-nivalis is similar to E. sakishimaense Otani (1976)
although the latter has larger basidiospores (15-24 x 5-6 um vs. 10.2-13 x
2.5-3 um).
The second new species parasitizes young leaves of Gaultheria pyroloides
(subfamily Gaultherioideae of Ericaceae), causing leaf spots that are red on
the upper surface. The host plant belongs to the subfamily Gaultherioideae of
Ericaceae. There are one or two diseased parts on each leaf. Transverse sections
of the diseased leaf show hypertrophy of plant cells. Hyphae protrude between
epidermal cells, forming a continuous thick layer on the lower surfaces of the
leaves at maturity. There are 2-4 sterigmata per basidium.
Exobasidium pyroloides ZhenYing Li & L. Guo, sp. nov. Figs. 2, 8-10
MycoBank MB 511911
Hymenium album, hypophyllum. Basidia cylindrica, 3.5-6 um lata, hyalina, terminaliter
2-4 sterigmatibus (2-)3-5(-6) x 1-1.5(-2) um praedita. Basidiosporae cylindricae vel
obovoideae, (7-)9-13 x 3-4(-5) um, hyalinae, laeves, primo continuae, dein 1(-2)-
septatae.
Hymenium white, hypophyllous. Basidia cylindrical, 3.5-6 tm wide, hyaline,
with 2-4 sterigmata. Sterigmata conical, (2—)3-5(-6) x 1-1.5(-2) um.
Basidiospores cylindrical or obovoid, (7—)9-13 x 3-4(-5) um, hyaline, smooth,
at first continuous, then 1(—2)-septate.
—
EE
Exobasidium spp. nov. (China) ... 333
10 um
Fig. 2. Line drawings of basidia, sterigmata and basidiospores of
Exobasidium pyroloides on Gaultheria pyroloides (HMAS 183432, holotype).
SPECIMEN EXAMINED—On Gaultheria pyroloides Hook.f. & Thomson ex Mig. (Ericaceae),
Yunnan: Tengchong, Xiaodifang, alt. 2180 m, 19 IX 2005, Z.Y. Li, L. Guo & N. Liu 201,
HMAS 183432 (holotype).
Exobasidium pyroloides is similar to E. gaultheriae Sawada on Gaultheria cuneata
Bean from which it mainly differs in having 2-4 sterigmata and forming leaf
spots; E. gaultheriae has 3-6 sterigmata and forms galls.
A third species of Exobasidium is a new Chinese record. This species is parasitic
on young leaves of Rhododendron sp., causing convex or concave galls on the
upper surface. The diseased parts are ellipsoidal, usually 9-13 x 4-6.5 mm in
size. There are one or more diseased parts on each leaf. Transverse sections of
the diseased leaf show that there is hypertrophy of plant cells. Hyphae protrude
between epidermal cells, forming a continuous layer on the lower surfaces of the
leaves at maturity. It is identified as Exobasidium cylindrosporum (Ezuka 1990),
which was described from Japan on Rhododendron macrosepalum Maxim..
Exobasidium cylindrosporum Ezuka, Trans. Mycol. Soc. Japan 31: 451, 1990.
Figs. 3, 11-13
10 um 3
i
Fig. 3. Exobasidium cylindrosporum on Rhododendron sp. (HMAS 183415).
A. Basidia, sterigmata and basidiospores. B. Basidiospore germination.
334 ... Li& Guo
Basidia cylindrical, 17-30 x 4-6(-8) um, hyaline, with 3-5(-6) sterigmata.
Sterigmata conical, (3.2-)4-5.5(-7) x 1.2-2 wm. Basidiospores cylindrical,
15-20.5 x (3.5-)4(-4.8) um, hyaline, smooth, at first continuous, with 3-8
septa when germinating, and slightly curved at the base. Short germ tubes
emerge from cells at both ends of the basidiospores and from near septa of
intermediate cells 24 h after dropping.
SPECIMEN EXAMINED—On Rhododendron sp. (Ericaceae), Jiangxi: Lushan, alt. 1080 m,
14 V 2007, Z.Y. Li & L. Guo 638, HMAS 183415.
Colonies on PDA grew slowly, to a maximum 10 mm diameter after 21 days
incubation at 25°C. The colony was yellow and corrugate on the surface,
composed of hyphae and conidia. Conidia linear, 9-20 x 1 um.
To date, 29 species of Exobasidium have been recorded in China (Sawada 1922,
Teng 1963, Tai 1979, Guo et al. 1991, Zang 1996, Li & Guo 2006a, b, 2008a, b)
including the three species in this paper.
Acknowledgements
The authors would like to express their deep thanks to Dr. Eric H.C. McKenzie and Robert
Bauer for serving as pre-submission reviewers, to Dr. Pennycook for nomenclatural
review, to Mr. Cao Ziyu (Institute of Botany, Chinese Academy of Sciences) for
identifying the host plants, to Mrs. Xie Jiayi and Liang Jingnan for assistance with SEM
photographs, and to Mrs. Zhu Xiangfei for inking in line drawings. This study was
supported by the National Natural Science Foundation of China (No. 30499340 and
No. 30670005).
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Guo L, Zhou YL, Li YB. 1991. Study of the genus Elaeodema and Exobasidium sawadae. Acta
Mycol. Sin. 10: 31-35.
Li ZY, Guo L. 2006a. A new species of Exobasidium (Exobasidiales) on Rhododendron from China.
Mycotaxon 96: 323-326. \
Li ZY, Guo L. 2006b. A new species and a new Chinese record of Exobasidium (Exobasidiales) from
China. Mycotaxon 97: 379-384.
Figs. 4-7. Exobasidium rhododendri-nivalis on Rhododendron nivale (HMAS 183431, holotype). Fig.
4. Symptoms. Fig. 5. Basidiospores and germ tubes as seen by LM. Fig. 6. Basidia and sterigmata as
seen by LM. Fig. 7. Basidia, sterigmata and basidiospores as seen by SEM. Figs. 8-10. Exobasidium
pyroloides on Gaultheria pyroloides (HMAS 183432, holotype). Fig. 8. Basidia and sterigmata as
seen by LM. Figs. 9. Basidiospores as seen by LM. Fig. 10. Basidia, sterigmata and basidiospores as
seen by SEM. Figs. 11-13. Exobasidium cylindrosporum on Rhododendron sp. (HMAS 183415). Fig.
11. Symptoms. Fig. 12. Basidium, sterigmata and basidiospores as seen by LM. Fig. 13. Basidium,
sterigmata and basidiospores as seen by SEM.
HV Det ‘Spot
Mag
Exobasidium spp. nov. (China) ... 335
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s
8
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rs
336 ... Li & Guo
Li ZY, Guo L. 2008a. Three new species and a new Chinese record of Exobasidium (Exobasidiales)
in China. Fung. Divers. (in the press).
Li ZY, Guo L. 2008b. Two new species of Exobasidium (Exobasidiales) from China. Mycotaxon
104: 331-336.
Otani Y. 1976. A new species of Exobasidium collected in Iriomote-island, Okinawa. Trans. Mycol.
Soc. Japan 17: 355-357.
Sawada K. 1922. Descriptive catalogue of the Formosan fungi. Part II. Dept. Agr. Govt. Res. Inst.
Formosa. Report 2. Exobasidiales. 106-110.
Tai FL. 1979. Sylloge Fungorum Sinicorum. Science Press, Beijing. 1-1527.
Teng SC. 1963. Fungi of China. Science Press, Beijing. 1-808.
Zang M. 1996. Fungi of the Hengduan Mountains. Science Press, Beijing. 1-598.
MY COTAXON
Volume 105, pp. 337-342 ea ca
| A new anamorphic fungus from Venezuela:
Pyriculariopsis formosa
TERESA ITURRIAGA
| titurri@usb.ve
| Departamento Biologia de Organismos, Universidad Simon Bolivar,
| Apartado 89000. Sartenejas, Baruta, Edo Miranda, Venezuela
ROBERTO FERNANDEZ
00-775@usb.ve
Departamento Biologia de Organismos, Universidad Simon Bolivar,
Apartado 89000. Sartenejas, Baruta, Edo Miranda, Venezuela
RAFAEL FEF, CASTANEDA RUIZ
rfcastaneda@inifat.co.cu
Instituto de Investigaciones Fundamentales en Agricultura Tropical Alejandro de Humboldt’ (INIFAT)
Calle 1 Esq. 2, Santiago de Las Vegas, C. Habana, Cuba, C.P. 17200
Davip W. MINTER
d.minter@cabi.org
Cabi Bioscience, Bakeham Lane
Egham, Surrey, TW20 9TY, United Kingdom
GABRIELA HEREDIA ABARCA
gabriela.heredia@inecol.edu.mx
Instituto de Ecologia, A.C., Km. 2.5 antigua carretera a Coatepec 325, Congregacion El Haya,
Xalapa, Veracruz, México. C.P. 91070
Abstract — Pyriculariopsis formosa anam. sp. nov. found on a decaying leaf of Clusia
rosea in ‘Cerro El Avila, Caracas, Venezuela, is described and ilustrated. This species
is characterized by erect, robust, unbranched conidiophores and obclavate, slightly
rostrate, 5- to 7-septate, smooth conidia with two dark brown cells and the other
conidial cells subhyaline. A key and illustrations of conidia of all Pyriculariopsis species
are provided.
Key words — conidial fungi, leaf litter, systematics, tropical rainforest
Introduction
During an expedition in 2007 to “Cerro El Avila” near Caracas, Venezuela, a
conspicuous anamorphic fungus from the genus Pyriculariopsis was collected.
338 ... Iturriaga & al.
Morphologically the fungus was distinctly different from previously described
species and is therefore described as a new species.
Materials and methods
Samples of plant material were placed in separate paper bags and taken to the
laboratory for examination of microfungi. Mounts were prepared in polyvinyl
alcohol-glycerol (8 gin 100 ml of water, plus 5 ml of glycerol) and measurements
made at a magnification of x 1000. Micrographs were obtained with a Leitz
Biomed microscope. |
Taxonomy
Pyriculariopsis formosa Rob. Fernandez, R. F. Castafieda & Iturr. anam. sp. nov.
MycoBank #: MB511740 Figs. 1-8
Ad omnibus speciebus Pyriculariopsis differt in conidiophoris robustis, eramosis, 86-230
um altis, 5-8 um crassis ad basim, atrobrunneis. Conidia obclavata, leviter rostrata, 5- ad
7-septata, laevia, sicca. 19-32 x 5-8 um, cum cellulis secunda et tertia atrobrunneis et
cetero dilute brunneis usque ad subhyalinis.
VENEZUELA, Caracas, “Cerro El Avila’, Parque Nacional El Avila, near “Lagunazo”
camping field, on decaying leaf of Clusia minor L. 10°33'1.9” N and 66°51°48.8" W, 2243
m alt, 20.X1.2007, coll. R. Fernandez. HOLOTYPE: VEN 395730.
ETyMo_ocy: Latin, formosus, meaning pretty and elegant.
CoLonIEs on the natural substratum effuse, pilose, epiphyllous, dark brown to
black. Mycelium mostly immersed composed of septate, branched, smooth-
walled, 2-4 um diam. brown hyphae. CONIDIOPHORES macronematous,
mononematous, robust, erect, straight or flexuous, 8 to 12-septate, dark brown,
smooth, 86-230 um tall, 5-8 um wide at the base. CONIDIOGENOUS CELLS
holoblastic, multilocal, integrated, terminal and intercalary, 6-24 x 4-6 um,
indeterminate, sympodially proliferating, with conspicuous, conical-truncated
ends, slightly melanized denticles (1-2 x 3 um); enteroblastic percurrent
proliferations rarely seen. Conrp1a solitary, obclavate, slightly rostrate at the
apex, 5 to 7-septate, acropleurogenous, smooth, 19-32 x 5-8 um, pale brown
to subhyaline, but with second and third cells dark brown, dry. TELEOMORPH:
unknown.
COMMENTSs: The genus Pyriculariopsis was introduced by Ellis (1971) for a single
species, P. parasitica M.B. Ellis. Matsushima (1975, 1989), Davydkina & Melnik
(1989), Castafieda & Kendrick (1990), Lai & Gao (1991) and Reddy et al (2002)
subsequently added seven species: Pyriculariopsis appendiculata Matsush., P.
miogae Matsush., P. breviphora Matsush., P. pleuroconidiophora Davydkina &
Melnik, P theobromae Castafieda & W.B. Kendr., P amomi (Z.D. Jiang & P.K.
Chi) X.H. Lai & H.L. Gao and P indica B.S. Reddy et al, but no taxonomic
account for the genus has been published. The main diagnostic features of
Pyriculariopsis formosa sp. nov. (Venezuela) ... 339
Figs. 1-7. Pyriculariopsis formosa from VEN 395730.
1-4. Conidia. 5-6. Detail of conidiogenous cells and conidia.
7. Conidiogenous cells and denticulate loci.
Bars = 10 um
340 ... Iturriaga & al.
8
Fig. 8. Pyriculariopsis formosa from VEN 395730.
Conidiophore, conidogenous cells and conidia. Bar = 10 um
Pyriculariopsis are the large, conico-truncate or cylindrical denticles produced
after holoblastic sympodial proliferations of the conidiogenous cells, obclavate,
subhyaline or hyaline, intermediate cells, and pale brown to brown conidia
which are released schizolytically. P pleuroconidiophora was transferred to the
genus Cacumisporium by Castafieda et al (2007). Published Pyriculariopsis
species cannot be confused with P_ formosa because of the difference in number
of conidial septa.
Pyriculariopsis formosa sp. nov. (Venezuela) ... 341
Figs. 9-15. Re-drawing of conidia of Pyriculariopsis species from literature.
9. PR amomi. 10. P. appendiculata. 11. P. breviphora. 12. P. indica.
13. P. miogae. 14. P. parasitica. 15. P theobromae.
Bars = 10 um
Key to accepted Pyrculariopsis species
1 (Onidsa SMOO tM awallede er me eae ie eee A ae oe een 2)
Conidia with second and sometimes third cells verrucose, 22-32 x 2.5-4 um,
2- to 3(4)-septate, fusiform , two central cells dark brown, end cells almost
colorless? cxportoeytS toh. wettnlstarrih-lo-eieadarts 7 P. theobromae (Fig.15)
2(1) Conidia 1-septate, cylindrical to fusiform, rostrate, 50-72 x 7.5-10 um,
pale brown, but with hyaline apical cell ............... P. breviphora* (Fig.11)
(Oras Ti WPA hes Vane t53 Clie peat Sees acmleeale ahead oa lrereie manent Ror. 2, R.ccrrinn Kena tir 3
Conidia 5- to 7-septate, 19-32 x 5-8 um, pale brown to subhyaline but
withisecond and thirdicellsidark brown M2, S76. ere. aa. tet. P. formosa
342 ... Iturriaga & al.
3(2) Conidiophores twisted near the base, conidia obpyriform to clavate,
20748:% 7-11 tm: pale brown © oboe nc ge eee P. amomi (Fig.9)
Conidiophores pot twisted 4c. se eos: ak Eat aes one a ene +
4(3) Conidia 1- to 2- septate, obclavate, 15-20 x 3-5 um, basal and suprabasal
cells subhyaline to pale brown, apical cell hyaline ........... P. indica (Fig.12)
Conidia 2- septate, rostrate, narrow obclavate, 16-28 x 3-4 um, basal and
suprabasal cells brown, apical cell hyaline ........... P. appendiculata (Fig. 10)
Conidiammore than 27septate 22.8 28. out Ph .4. 6 ee ee oe 5
5(4) Conidia 3-septate, obclavate, rostrate, 30-55 x 9-12 um, end cells hyaline,
intermediatercells pale browngs..2..2%. 4.44.04. -8- eaten P. parasitica (Fig.14)
Conidia 3-septate, broad or narrow fusiform, 45-60 x 6-8 um, end cells
hyaline; intermediate cells‘pale brown,2. 2s... .-). eee P. miogae (Fig.13)
* Described from pure culture on V8 Juice agar.
Acknowledgements
We are deeply indebted to Prof. Lori M. Carris (Washington State University) and Dr.
Marc Stadler (Univ. Bayreuth), for kindly reviewing the manuscript. We thank Ciencia y —
Tecnologia para el Desarrollo (CYTED RED-XILJ), the Cuban Ministry of Agriculture
for facilities as well as Universidad Simon Bolivar’s Decanato de Investigacion y
Desarrollo for RFCR funding to stay at this university. Part of the support for this work
came from the UK Darwin Initiative.Collecting in Venezuela has been possible through
permits obtained from INPARQUES (0721) and MARN (1-096), institutions to whom
we are indebted. The author RFCR thanks Drs. Pedro Crous, Uwe Braun, Ludmila
Marvanova, Cony Decock, Jerry A. Cooper, L.M. Carris, Antonio Hernandez-Gutiérrez
and Josep Cano for their generous and valuable assistance with literature not available
in our libraries.
Literature cited
Castaneda Ruiz RF, Kendrick WB. 1990. Conidial fungi from Cuba II. University of Waterloo
Biology Series. 33: 1-61.
Castafieda Ruiz RF, Heredia Abarca G, Arias RM, Saikawa M, Stadler M. 2007. Anamorphic fungi
from submerged plant material: Phaeomonilia pleiomorpha, P. corticola and Cacumisporium
pleuroconidiophorum. Mycotaxon 100: 327-336.
Davydkina TA, Mel'nik VA. 1989. Two new hyphomycetes from genera Cordana and Pyriculariopsis.
Mikol. Fitop. 23(2): 110-113. [in Russian]
Ellis MB. 1971. Dematiaceous hyphomycetes. Commonwealth Mycological Institute, Kew, Surrey.
Lai XH, Gao HL. 1991. A new combination of Pyriculariopsis. Acta Mycologica Sinica.10: 79-80.
Matsushima T. 1975. Icones microfungorum a Matsushima lectorum. Osaka, published by the
author.
Matsushima T. 1989. Matsushima Mycological Memoirs. 6: 36. Kobe published by the author.
Reddy BS, Manoharachary C, Agarwal DK, Rao V. 2002. Two new hyphomycetes from India.
Indian Phytopath. 55: 213-215.
MYCOTAXON
Volume 105, pp. 343-348 July-September 2008
Wood-rotting fungi in eastern China 2.
A new species of Fomitiporia (Basidiomycota) from
Wanmulin Nature Reserve, Fujian Province
BAao-Kai Cul
baokaicui@yahoo.com.cn
: Institute of Microbiology, Beijing Forestry University
| Beijing 100083, China
Yu-CHENG Dal
*Corresponding author, yuachengd@yahoo.com
Institute of Microbiology, Beijing Forestry University
Beijing 100083, China
Institute of Applied Ecology, Chinese Academy of Sciences
Shenyang 110016, China
Abstract — Fomitiporia ellipsoidea sp. nov., collected from Wanmulin Nature Reserve,
Fujian Province, eastern China, is described and illustrated. It has a perennial growth
habit, resupinate basidiocarp, yellowish brown to rust brown pore surface, dimitic
hyphal structure with simple septa, abundant hymenial setae, dextrinoid, cyanophilous
and thick-walled basidiospores. It is distinguished from the other species in the genus
by having hooked hymenial setae and ellipsoid to broadly ellipsoid basidiospores.
Key words — Hymenochaetaceae, lignicolous and poroid fungi, taxonomy
Introduction
Fomitiporia Murrill was established by Murrill (1907), and today the genus is
often conceived as the Phellinus robustus complex. Species of Fomitiporia have
hyaline, thick-walled, dextrinoid and cyanophilous basidiospores, and these
characters distinguish the genus from Phellinus sensu lato. Recent phylogenetic
studies have demonstrated that Fomitiporia is a distinct group in the family
Hymenochaetaceae (Decock et al. 2005, Fischer 1996, 2002, Wagner & Fischer
2001, 2002).
Nearly 20 taxa have been transferred to or described in the genus (Dai & Cui
2005, Dai et al. 2001, 2008, Dai & Zang 2002, Decock et al. 2005, Fiasson &
Niemela 1984, Fischer 2002, Fischer & Binder 2004, Fischer et al. 2005), and
344 ... Cui & Dai
10 species of Fomitiporia have previously been reported from China (Cui et al.
2007, Dai 1999, Dai & Cui 2005, Dai et al. 2001, 2008, Dai & Niemela 2006, Dai
& Zang 2002).
During the study of wood-rotting fungi from Wanmulin Nature Reserve in Fujian
Province, eastern China, five specimens of an unknown taxon were collected,
characterized by perennial, resupinate basidiocarps, dimitic hyphal structure
with simple septa, abundant hymenial setae, and basidiospores ellipsoid to
broadly ellipsoid, hyaline, thick-walled, dextrinoid and cyanophilous. These
characters indicate the genus Fomitiporia, but with no existing species name
for this taxon, in this paper we describe it as new.
Materials and methods
The studied specimens are deposited at the herbarium of Institute of Applied
Ecology, Chinese Academy of Sciences (IFP), the herbarium of Beijing
Forestry University (BJFC), and the herbarium of the botanical museum of the
University of Helsinki (H). The microscopic procedure follows Cui et al. (2007).
In presenting the variation in the size of the spores, 5% of measurements were
excluded from each end of the range, and are given in parentheses. In the text
the following abbreviations are used: IKI = Melzer’s reagent, IKI- = negative
in Melzer’s reagent, KOH = 5% potassium hydroxide, CB = Cotton Blue, CB+
= cyanophilous, CB- = acyanophilous, L = mean spore length (arithmetic
mean of all spores), W = mean spore width (arithmetic mean of all spores),
Q = variation in the L/W ratios between the specimens studied, n = number
of spores measured from given number of specimens. Sections were studied at
magnification up to x1000 using a Nikon Eclipse E600 microscope and phase
contrast illumination. Drawings were made with the aid of a drawing tube.
Special colour terms are from Petersen (1996) and Anonymous (1969).
Description
Fomitiporia ellipsoidea B.K. Cui & Y.C. Dai, sp. nov. Fig.1
MycoBank MB 512012
Carpophorum perenne, resupinatum, contextum fulvum. Facies pororum fulva vel
ferrugineum; pori rotundi vel sinuolati, 5-8 per mm. Systema hypharum dimiticum,
hyphae generatoriae septatae, efibulatae. Sporae ellipsoideae, crassitunicatae, dextrinoidae,
CB+, 4.9-6 x 3.9-4.8 um.
Type. — China. Fujian Province, Jianou County, Wanmulin Nature Reserve, on fallen
angiosperm trunk, 31.VIII.2006 Cui 4285 (holotype in IFP, isotypes in BJFC and H).
EtyMo.Locy — Ellipsoidea (Lat.): referring to ellipsoid basidiospores.
FRUITBODY — Basidiocarps perennial, resupinate, inseparable, woody hard
when fresh, without odour or taste, consistency woody hard when dry, up to 30
Fomitiporia ellipsoidea sp. nov. (China) ... 345
Fig. 1. Microscopic structures of Fomitiporia ellipsoidea (drawn from the holotype).
—a: Basidiospores. —b: Basidia and basidioles. —c: Hymenial setae.
—d: Hyphae from trama. —e: Hyphae from subiculum.
346 ... Cui & Dai
cm or more in longest dimension, 20 cm wide, ca. 8 mm thick at centre; margin
yellowish to yellowish brown, up to 2 mm wide. Pore surface yellowish brown
to rust brown, shining; pores circular to sinuous at oblique surface, 5-8 per
mm; dissepiments slightly thick, entire. Context yellowish brown, woody hard,
very thin, less than 0.5 mm thick. Tubes concolourous with the pore surface,
woody hard, up to 8 mm long, distinctly stratified.
HYPHAL STRUCTURE — Hyphal system dimitic; all septa without clamp
connections; skeletal hyphae IKI-, CB-; tissue darkening but otherwise
unchanged in KOH. |
SUBICULUM — Tissue dominated by skeletal hyphae; generative hyphae hyaline,
thin- to fairly thick-walled, occasionally branched and frequently simple
septate, 2-3 um in diam.; skeletal hyphae golden brown to rust brown, thick-
walled with a narrow to wide lumen, unbranched, interwoven, more or less -
agglutinated, 2-3.6 um diam.
TuBes — Tramal hyphae dominated by skeletal hyphae; generative hyphae
hyaline, thin- to slightly thick-walled, occasionally branched and frequently
simple septate, 1.5-2.6 um diam.; skeletal hyphae yellowish brown to rust
brown, thick-walled to subsolid, unbranched, interwoven, slightly agglutinated,
1.8-3.4 um diam. Subhymenium indistinct. Hymenial setae present, usually
ventricose and hooked, yellowish brown, thick-walled, 20-30 x 10-14 um;
cystidia and cystidioles absent; basidia barrel-shaped to subglobose, with four
sterigmata and a simple septum at the base, 8-12 x 6-7 um; basidioles in shape
similar to basidia, but slightly smaller; big rhomboid crystals abundant in
hymenia and trama.
Spores — Basidiospores ellipsoid to broadly ellipsoid, hyaline, slightly thick-
walled, smooth, weakly dextrinoid in Melzer’s reagent, distinctly CB+, (4.5-)
4.9-6(-6.1) x (3.5-)3.9-4.8(-5) um, L = 5.26 um, W = 4.14 um, Q = 1.22-1.32
(n=90/3).
ADDITIONAL SPECIMENS (PARATYPES) EXAMINED. — China. Fujian Province, Jianou
County, Wanmulin Nature Reserve, on fallen angiosperm trunk, 30.VIII.2006 Cui 4234
(IFP, BJFC); 31. VIII.2006 Cui 4270, 4277 & 4289 (IFP, BJFC).
TYPE OF ROT — White rot.
REMARKS — Fomitiporia ellipsoidea has a very distinctive morphology with
its perennial growth habit, resupinate basidiocarp, hooked hymenial setae,
ellipsoid, weakly dextrinoid and cyanophilous basidiospores. It is easily
recognized under the microscope because of its hooked setae and ellipsoid
basidiospores.
The dextrinoid reaction of basidiospores of E ellipsoidea is not as strong as in
most species of Fomitiporia, and basidiospore shape is also not typical of the
Fomitiporia ellipsoidea sp. nov. (China) ... 347
genus. All other features, however, fit the genus well and on that basis we treat
it here under Fomitiporia.
Five species in Fomitiporia, E bannaensis Y.C. Dai, F. pseudopunctata
(A. David et al.) Fiasson, FE sonorae (Gilb.) Y.C. Dai, F. sublaevigata (Cleland
& Rodway) Y.C. Dai, and E tenuis Decock et al., have resupinate basidiocarps
and hymenial setae (Dai 1999, Dai et al. 2001, Decock et al. 2005), but all these
five species have straight hymenial setae, and their basidiospores are basically
subglobose to globose.
Phellinus caribaeo-quercicola Decock & S. Herrera was recently described from
Cuba (Decock et al. 2006), and this species shares with Fomitiporia ellipsoidea
the hooked hymenial setae and ellipsoid to broadly ellipsoid basidiospores.
However, it differs from F ellipsoidea in having resupinate to effused basidiocarps
and hyaline to yellowish, nondextrinoid basidiospores. Moreover, it is only
known from Quercus cubana from tropical America (Decock et al. 2006).
Phellinus uncinatus Rajchenb. has hooked hymenial setae, thick-walled and
dextrinoid basidiospores (Decock et al. 2006, Rajchenberg 1987), but its
basidiospores are distinctly subglobose to globose, and bigger than those of
E ellipsoidea (5.5-7 x 5-6.5 um, Ryvarden 2004). Furthermore, P. uncinatus
grows on bamboo exclusively in tropical America (Decock et al. 2005).
Acknowledgements
Special thanks are due to Dr. Hai-Sheng Yuan (Shenyang, China) for technical assistance
with the drawings. We express our gratitude to Prof. Teuvo Ahti (H, Finland) for revising
the Latin description and improving the manuscript, and to Drs. Peter Buchanan
(Landcare Research, New Zealand) and Urmas Koljalg (University of Tartu, Estonia)
who reviewed the manuscript. The research was financed by Beijing Forestry University
(Project No. BLX2007014), the National Natural Science Foundation of China (Project
No. 30425042) and the Chinese Academy of Sciences (Project on taxonomy).
Literature cited
Anonymous. 1969. Flora of British fungi. Colour identification chart. Her Majesty’s Stationery
Office, London.
Cui BK, Dai YC, Decock C. 2007. A new species of Perenniporia (Basidiomycota, Aphyllophorales)
from eastern China. Mycotaxon 99: 175-180.
Cui BK, Dai YC, Knudsen H. 2007. Two species of Hymenochaetaceae (Basidiomycota,
Aphyllophorales) new to China. Mycosystema 26: 144-147,
Dai YC. 1999. Phellinus sensu lato (Aphyllophorales, Hymenochaetaceae) in East Asia. Acta Bot.
Fennica 166: 1-115.
Dai YC, Cui BK. 2005. Two new species of the Hymenochaetaceae from eastern China. Mycotaxon
94: 341-347.
348 ... Cui & Dai
Dai YC, Cui BK, Decock C. 2008. A new species of Fomitiporia (Hymenochaetaceae, Basidiomycota)
from China based on morphological and molecular characters. Mycological Research, 112:
375-380.
Dai YC, Niemela T. 2006. Hymenochaetaceae in China: hydnoid, stereoid and annual poroid genera,
plus additions to Phellinus. Acta Bot. Fennica 179: 1-78.
Dai YC, Zang M. 2002. Fomotiporia tibetica, a new species of Hymenochaetaceae (Basidiomycota)
from China. Mycotaxon 83: 217-222.
Dai YC, Zhang XQ, Zhou TX. 2001. New and noteworthy species of Hymenochaetaceae from
China. Mycosystema 20: 16-21.
Decock C, Bitew A, Castillo G. 2005. Fomitiporia tenuis and Fomitiporia aethiopica (Basidiomycetes,
Hymenochaetales), two undescribed species from the Ethiopian highlands: taxonomy and
phylogeny. Mycologia 97: 121-129.
Decock C, Herrera FS, Robledo G, Castillo G. 2006. Phellinus caribaeo-quercicolus sp. nov., parasitic
on Quercus cubana, taxonomy and preliminary phylogeny. Mycologia 98: 265-274.
Fiasson JL, Niemela T. 1984. The Hymenochaetales: a revision of the European poroid taxa.
Karstenia 24: 14-28.
Fischer M. 1996. On the species complexes within Phellinus: Fomitiporia revisited. Mycological
Research 100: 1459-1467.
Fischer M. 2002. A new wood-decaying Basidiomycetes species associated with esca of grapevine:
Fomitiporia mediterranea (Hymenochaetales). Mycological Progress 1: 315-324.
Fischer M, Binder M. 2004. Species recognition, geographic distribution, and host-pathogen
relationships: a case study in a group of lignicolous Basidiomycetes, Phellinus s.1. Mycologia
967 799-31
Fischer M, Edwards J, Cunnington JH, Pascoe IG. 2005. Basidiomycetous pathogens on grapevine:
a new species from Australia - Fomitiporia australiensis. Mycotaxon 92: 85-96.
Murrill WA. 1907. Polyporaceae. — North American Flora 9: 1-131.
Petersen JH. 1996. Farvekort. The Danish Mycological Society’s colour-chart. Foreningen til
Svampekundskabens Fremme, Greve.
Rajchenberg M. 1987. New South American polypores. Mycotaxon 28: 111-118.
Ryvarden L. 2004. Neotropical polypores 1. Syn. Fungorum 19: 1-229.
Wagner T, Fischer M. 2001. Natural groups and a revised system for the European poroid
Hymenochaetales (Basidiomycota) supported by nLSU rDNA sequence data. Mycological
Research 105: 773-782. |
Wagner T, Fischer M. 2002. Proceedings towards a natural classification of the worldwide taxa
Phellinus s. |. and Inonotus s. |., and phylogenetic relationships of allied genera. Mycologia 94:
998-1016.
MYCOTAXON
Volume 105, pp. 349-354 July-September 2008
Notes on Elmerina and Protomerulius (Basidiomycota)
YU-LIAN WEI
weiyulianer@hotmail.com
Institute of Applied Ecology, Chinese Academy of Sciences
Shenyang 110016, China
Yu-GHENGIDAT
*Corresponding author, yuchengd@yahoo.com
Institute of Applied Ecology, Chinese Academy of Sciences
Shenyang 110016, China
Abstract — Polypores with longitudinally septate basidia are briefly summarized, and a
key for 11 species is supplied. Protomerulius efibulatus is described as new from China.
It is characterized by resupinate basidiocarps, cream to pinkish buff pores, generative
hyphae without clamp connections, and ellipsoid basidiospores. It was found on rotten
angiosperm wood in tropical forests of southern China.
Key words — lignicolous fungi, poroid fungi, taxonomy
Introduction
Two polypore genera, Elmerina Bres. and Protomerulius Moller, are
macroscopically similar to polypores, having poroid and corky basidiocarps
when fresh. However, microscopically species of these two genera have
longitudinally septate basidia. Until now, Protomerulius caryae (Schwein.)
Ryvarden, P africanus (Ryvarden) Ryvarden and P. brasiliensis Moller have
been recorded in the genus Protomerulius; the first-mentioned species occurs
commonly in the northern hemisphere and also in Australasia (Gilbertson
& Ryvarden 1986, Ryvarden & Gilbertson 1993, Nufiez & Ryvarden 2001).
Protomerulius africanus and P. brasiliensis were reported from African and
South America (Ryvarden & Johansen 1980, Bandoniet al. 1982). Seven species
of Elmerina have previously been reported: E. borneensis (Jiilich) D.A. Reid,
E. cladophora (Berk.) Bres., E. dimidiata (A. David) D.A. Reid, E. hexagonoides
(A. David & Jag.) Nunez, E. holophaea (Pat.) Parmasto, E. substuppea (Berk.
& Cooke) T. Hatt. and E. unguliformis Corner (Parmasto 1984, Corner 1989,
Reid 1992, Nufiez 1998, Hattori 2003).
350 ... Wei & Dai
During an investigation of wood-inhabiting fungi in Hainan Province, tropical
China, two special specimens were collected with distinctly poroid, resupinate
basidiocarps, septate probasidia and longitudinally septate epibasidia. These
characters suggest the species belongs in Protomerulius (Nunez 1998), but no
existing name is available; we therefore describe them herein as a new species.
In addition, based mostly on published descriptions, a key to the two polypore
genera, Elmerina and Protomerulius, is supplied.
Materials and methods
The studied specimens are deposited at the herbarium of Institute of Applied
Ecology, Chinese Academy of Sciences (IFP), and the Herbarium of Beijing
Forestry University (BJFC). The microscopic procedure follows Yuan et al.
(2007). In presenting the variation in the size of the spores, 5% of measurements
were excluded from each end of the range, and are given in parentheses.
In the text the following abbreviations are used: IKI = Melzer’s reagent, IKI- =
negative in Melzer’s reagent, KOH = 5% potassium hydroxide, CB = Cotton
Blue, CB+ = cyanophilous, CB- = acyanophilous, L = mean spore length ~
(arithmetic mean of all spores), W = mean spore width (arithmetic mean of
all spores), Q = variation in the L/W ratios between the specimens studied, n
= number of spores measured from given number of specimens. Sections were
studied at magnification up to x1000 using a Nikon Eclipse E600 microscope
and phase contrast illumination. Drawings were made with the aid of a drawing
tube. Special colour terms are from Petersen (1996) and Anonymous (1969).
Description
Protomerulius efibulatus Y.C. Dai & Y.L. Wei, sp. nov. Fig.1
MycoBAankK MB 512016
Carpophorum annuum, resupinatum, facies pororum cremea vel bubalina; pori rotundi,
6-8 per mm. Systema hypharum dimiticum, hyphae generatoriae septatae, efibulatae;
hyphae skeletales subiculi 2.5-3 um in diam. Basidia longitudinaliter vel oblique septata,
sporae ellipsoideae, hyalinae, IKI-, CB-, 4.3-5.5 x 2.5-3.3 um.
Type. — China. Hainan Province, Lingshui County, Diaoluoshan Nature Reserve, on
fallen angiosperm trunk, 21.X1.2007 Dai 9378 (holotype in IFP).
ErymMo.Locy — Efibulatus (Lat.): referring to the simple septa of generative hyphae.
FRUITBODY — Basidiocarps annual, resupinate, difficult to separate from the
substrate, soft corky when fresh, becoming corky upon drying, up to 4.5 cm
long, 2.5 cm wide and 0.5 mm thick. Pore surface cream to pale buff, colour
changed to pale brownish when bruised, almost unchanged upon drying; sterile
margin very narrow; pores round, 6-8 per mm; dissepiments thin, entire when
juvenile, lacerate with age. Context cream, corky, very thin, less than 0.2 mm
thick. Tube layer concolorous with pore surface, up to 0.5 mm long.
Protomerulius efibulatus sp. nov. (China)... 351
a,
Y
908.
atthe A
comme,
6
uw
be >
ane 2 rots fy Cade *
ten... An by
eae 2 b
X ”,
2S
ES
——— WSS
0
gi
=
>
qi CF
=i = Vi me STE) LS
Aa SK
Y/ poe YE SV
= cee 10 pm
Fig. 1. Microscopic structures of Protomerulius efibulatus (drawn from the holotype).
—a: Basidiospores. —b: Epibasidia and probasidia.
—c: Hyphae from trama. —d: Hyphae from subiculum.
352 ... Wei & Dai
HyPHAL STRUCTURE — Hyphal system dimitic; generative hyphae bearing
simple septa; skeletal hyphae dominant, thick-walled, rarely branched, IKI-,
CB+; tissues unchanged in KOH.
SUBICULUM — Generative hyphae scanty, hyaline, thin-walled, occasionally
branched, 2-3 um diam.; skeletal hyphae dominant, hyaline, thick-walled with
a narrow lumen to subsolid, flexuous, rarely branched, interwoven, 2.5-3 um
diam.
TuBes — Generative hyphae scanty, hyaline, thin-walled, rarely branched,
2-3 um diam.; skeletal hyphae thick-walled with a narrow lumen to subsolid,
flexuous, sometimes encrusted with granular crystals, rarely branched, tightly
interwoven, 2-3.5 um diam. Cystidia and cystidioles absent. Probasidia
pyriform to subglobose, thin-walled, longitudinally or obliquely septate, 6-8.5
x 5.3-7.5 tm; epibasidia barrel-shaped, thin-walled, longitudinally septate,
with four sterigmata and a simple septum at the base, 8-10 x 6-7 um.
SpoRES — Basidiospores ellipsoid, hyaline, thin-walled, smooth, IKI-,
CB-, (4-)4.3-5.5(-6) x (2.1-)2.5-3.3(-3.6) um, L = 4.98 um, W = 2.97 um,
Q = 1.63-1.73 (n=66/2).
ADDITIONAL SPECIMEN (PARATYPE) EXAMINED. — China. Hainan Province, Ledong
County, Jianfengling Nature Reserve, on rotten angiosperm wood, 19.XI.2007 Dai 9322
(IFR):
Discussion
Previously three polypores, Protomerulius brasiliensis, P. caryae and Elmerina
borneensis, shared characters of a poroid hymenophore, resupinate basidiocarps
and longitudinally septate basidia. Protomerulius brasiliensis is distinguished
from P. efibulatus by its large pores (1-2 per mm) and big basidiospores
(6-6.5 x 4-5 um, Bandoni et al. 1982). Protomerulius caryae differs from P.
efibulatus by its greyish pores, allantoid basidiospores (4.9-6.2 x 2.2-2.8 um,
Niemela 2005), and clamp connections on its generative hyphae. In addition, P.
caryae occurs in temperate forests (Gilbertson & Ryvarden 1986, Ryvarden &
Gilbertson 1993, Nufez & Ryvarden 2001).
Elmerina borneensis is different from P. efibulatus by its bigger pores (1-2 per
mm) and bigger basidiospores (8-10 x 5-6 um, Nufez 1998). In addition, its
probasidia are clavate and aseptate.
Protomerulius and Elmerina were included previously in Tremellales, but recent.
phylogenetic studies showed both genera were closely related to Auriculariales
rather than Tremellales (WeiB & Oberwinkler 2001, Larsson et al. 2004). We
did not make molecular study to indicate the phylogenetic relationships of the
new species and other taxa in these two genera, such work will be done in the
coming years.
Protomerulius efibulatus sp. nov. (China)... 353
Key to species of E/merina and Protomerulius
ie Obasidia pyritormytosubeloboseusepiate . &. ee. Act a: 2 (Protomerulius)
MEEODASIOIAC AV ALCCASCD IG Lee guar mE Mane ee wep aes. gioatvccile peacncsicnlawsavrecrecsuels 5 (Elmerina)
puGenerative hyphaewith simplesepta.n. mote, Ones. Wahl, L000 LT Jone P. efibulatus
mmacierative Nypnac With, clampicannections awe. ss. ves oe ene eee Oe ee 3
Bm Aesj— 4 per (il DaslclocspOres allantOlGmee swine 6 ee fas oe ee seen ic P. caryae
Serores 1-2 per mm, basidigspores ellipsoid’ OMOVal. “rite “eit. areca ascnek wilde: 4
HGlococystidia presents, basidiosporesiovales. delist! oc). adeonel. .2ee P. brasiliensis
4, Gloeocystidia absent; basidiospores ellipsoid......................... P. africanus
Eepasidiocal ps TeSUpIiate = eat eee ER ALONE POOR en. 5 E. borneensis
BMD AsICIOCALPS DILCALCteneme tia, Wears Ss Maree eR rar Eee Pe eae CNS lot et ake pairs 6
6. Hymenophore lamellate to almost daedalioid ....................... E. holophaea
Darmenophorepporoid tompartlplamellatesaaaake Autos set. Paige. tarsi eewlawe, .. ji
Rm a SICOCALDS CII G1at Cage, eee eee ee i i tL, allt 8
Pm OSICLOCALDS SUDSUD Ita leg Bat Eh Pere Br SERED ra te ts a
Bmeoressiially Goci Peraniie st wey Rite Nee wees eee ct ar E. dimidiata
PEBOLESIDID al OLIN Wid Cats eee eRe re oe ee. E. hexagonoides
Pabasidiospores sinaller Sal Inlenothwr: sameeren lessee eels ele E. substuppea
ee asIGIOspOLes Digvet es .o WIT LENO Ul ge a) eres tee Or Sie Sore a ay ae ee 10
10. Hyphal pegs present; skeletal hyphae dominant.................... E. cladophora
10. Hyphal pegs absent; generative hyphae dominant................ E. unguliformis
Acknowledgements
We express our gratitude to Prof. Teuvo Ahti (University of Helsinki, Finland) for
helping with the preparation of the Latin description, and to Drs. Peter Buchanan
(Landcare Research, New Zealand) and Urmas KOljalg (University of Tartu, Estonia)
who reviewed the manuscript. The project is supported by the National Natural Science
Foundation of China (Project No. 30500003) and the Chinese Academy of Science
(Project on taxonomy).
Literature cited
Bandoni R, Oberwinkler FE, Wells K. 1982. On the poroid genera of the Tremellaceae. Canadian J.
Bot. 60: 998-1003.
Corner EJH. 1989. Ad Polyporaceas 5. Beih. Nova Hedwigia 96: 1-218.
Gilbertson RL, Ryvarden L. 1986. North American polypores 1. Abortiporus — Lindtneria.
Fungiflora, Oslo. 1-433 pp.
Hattori T. 2003. Type studies of the polypores described by E.J.H. Corner from Asia and West
Pacific areas 5. Species described in Tyromyces (2). Mycoscience 44: 265-276.
Larsson K-H, Larsson E, Kéljalg U. 2004. High phylogenetic diversity among corticioid
homobasidiomycetes. Mycological Research 108: 983-1002.
354 ... Wei & Dai
Niemela T. 2005. Polypores, lignicolous fungi. Norrlinia 13: 1-320. (in Finnish, with English
summary).
Nuifiez M. 1998. The genus Elmerina (Heterobasidiomycetes) in Japan. Folia Cryptog. Estonica 33:
99-101.
Nunez M, Ryvarden L. 2001. East Asian polypores 2. Polyporaceae s. lato. — Synopsis Fungorum
14: 170-522.
Parmasto E. 1984. The genus Elmerina (Polyporaceae s. str.). Nova Hedwigia 39: 101-116.
Reid DA. 1992. The genus Elmerina (Tremellales) with accounts of two species from Queensland,
Australia. Persoonia 14: 465-474.
Petersen JH. 1996. Farvekort. The Danish Mycological Society’s colour-chart. Foreningen til
Svampekundskabens Fremme, Greve. 6 unnumbered pp.
Ryvarden L, Gilbertson RL. 1993. European polypores 2. Synopsis Fungorum 7: 388-743.
Ryvarden L, Johansen I. 1980. A preliminary polypore flora of East Africa. Fungiflora, Oslo. 636
PP:
WeiB M, Oberwinkler, F. 2001. Phylogenetic relationships in Auriculariales and related groups-
hyphotheses derived from nuclear ribosomal DNA sequences. Mycological Research 105:
403-415.
Yuan HS, Li J, Huang MY, Dai YC. 2006. Antrodiella stipitata sp. nov. from Heilongjiang Province,
northeastern China, and a critical checklist of polypores from the area. Cryptogamie Mycologie
27: 21-29.
CT
MYCOTAXON
Volume 105, pp. 355-377 July-September 2008
Lepiota (Agaricaceae, Basidiomycota)
species diversity in Israel
ANUSH KOSAKYAN', MARYNA DIDUKH?’, YAIR UR3,
SOLOMON P. WASSER *” & EVIATAR NEVo!
akosakyan@yahoo.com
‘Institute of Evolution, and Department of Evolutionary and Environmental Biology
Faculty of Science, University of Haifa, Mt. Carmel, Haifa 31905, Israel
°M.G. Kholodny Institute of Botany, National Academy of Sciences of Ukraine
2 Tereshchenkivska St., Kiev 01601, Ukraine
3Amnum, P.O. Box 106, 12392, Israel
Abstract — Species diversity for Lepiota in Israel is presented. The 18 Lepiota species
recorded from Israel include L. apatelia, L. brunneoincarnata, L. brunneolilacea,
L. castanea, L. clypeolaria, L. cortinarius, L. echinella, L. erminea, L. forquignonii,
L. helveola, L. ignivolvata, L. lilacea, L. locquinii, L. micropholis, L. oreadiformis, L.
pallida, L. subgracilis, and L. subincarnata. Eight species —L. cortinarius, L. echinella,
L. forquignonii, L. helveola, L. ignivolvata, L. locquinii, L. oreadiformis, and L. pallida
— are reported for the first time for Israel. Israeli locations and collection dates, habitat
data, general distribution, and taxonomic remarks are provided for all species. Detailed
macro- and micromorphological descriptions and illustrations are given for the eight
species newly recorded from Israel.
Key words — Asia, biodiversity, ecology, Lepioteae, taxonomy
Introduction
Lepiota is one of the most problematic genera in the Agaricaceae. Traditional
taxonomic classification in the genus is based on a very limited number of traits,
such as spore shape and size, pileipellis structure, microchemical reactivity, and
fruitbody habitus. An overview of current Lepiota systematics reveals a lack of
taxonomic consensus regarding which infrageneric taxa to recognize and which
traits are of infrageneric significance (Candusso & Lanzoni 1990, Wasser 1980,
Migliozzi 1996). Molecular datasets suggest a need for total reorganization of
current Lepiota systematics, which conflicts with phylogenetic relationships
inferred from molecular analyses (Johnson & Vilgalys 1998, Johnson 1999,
356 ... Kosakyan & al.
Vellinga 2003, 2004a). Nonetheless, we feel that extensive study of known taxa,
especially those described from the tropics, is required prior to any serious
taxonomic changes.
To date, approximately 500 species are included in the genus Lepiota (Candusso
& Lanzoni 1990, Guzman & Guzman-Davalos 1992, Kirk et al. 2001, Vellinga
2004b). The genus is almost cosmopolitan, with most species tropical and
subtropical in distribution (Singer 1986). Nonetheless, although Vellinga
(2004b) notes that Leucoagaricus and Leucocoprinus are more species rich in
the tropics than in temperate regions, Lepiota does occur in relatively cold areas
(high latitudes, high altitudes).
The genus Lepiota is not well studied in Israel. Fragmentary data on some
Lepiota species is provided in papers devoted to Agaricales from Israel
(Avizohar-Hershenzon 1968, Binyamini 1975, 1976, 1984; Didukh et al. 2002,
Kosakyan et al. 2008). Prior to our current research only ten Lepiota species
were reported for Israel: L. apatelia, L. brunneoincarnata, L. brunneolilacea,
L. clypeolaria, L. castanea, L. erminea, L. lilacea, L. micropholis, L. subgracilis,
and L. subincarnata. Our investigation now adds eight Lepiota species to the —
known Israeli mycobiota: L. cortinarius, L. echinella, L. forquignonii, L. helveola,
L. ignivolvata, L. locquinii, L. oreadiformis, L. pallida. Of these, L. apatelia, L.
brunneolilacea, L. cortinarius, L. echinella, L. ignivolvata, and L. pallida, are new
reports for Asia.
Material and methods
The species-richness research on Lepiota species in Israel is based on (1) our
fieldwork during 1995-2007 fruiting seasons; (2) a study of extensive material
at the Tel Aviv University herbarium (TELA) in April 1991, and (3) a review
of the published literature on the Agaricales of Israel. The material collected is
kept at the Herbarium of the Institute of Evolution, University of Haifa (HAI,
Israel).
Microscopic characteristics were observed using a Carl Zeiss-amplival
microscope. Fungal material was mounted on a microscope slide and examined
in water using a light/dark field microscope with or without phase contrast
at x20, x40, and x100 (oil immersion). For statistical calculations, 30 spores
were measured for every preparation. Micromorphological characteristics of
our specimens were observed using Melzer’s reagent and Cresyl blue. General
species distributions are given according to Wasser (1980), Moser (1983),
Candusso & Lanzoni (1990), Guzman & Guzman- Davalos (1992), and Vellinga
(2001, 2004b). Distribution (Fic. 1) of species in Israel is given according to
natural regions of Israel (Feinbrun-Dothan & Danin 1998).
Lepiota species diversity in Israel ... 357
FicurE 1. Natural regions of Israel: AP - Akko
Plain; AV - Arava Valley; BS - Beit Shean
Valley; CC - Carmel Coast; CG - Coast Galilee;
CM - Carmel Mount; CN - Central Negev;
DS - Dead Sea Area; EP - Esdraelon (Yizreel)
Plain; GH - Golan Heights; GM - Gilboa
Mount; HE - Hermon Mount; HP - Hula Plain;
JD - Judean Desert; JM — Judean Mts.; LG - Lower
Galilee; LJ - Lower Jordan Valley; NN - Northern
Negev; PP - Philistean Plain; SA - Samaria;
SH - Shefela; SN - South Negev; SP - Sharon
Plain; UG - Upper Galilee; UJ - Upper Jordan
Valley; WN - Western Negev.
Taxonomy
The description of species diversity of genus Lepiota of Israel is presented below,
including eight species new to Israel and a number new to Asia or Near East.
Genus Lepiota
(Pers.) Gray, A Natural arrangement of British plants 1: 601 (1821).
Lepiota differs from the closely related genus Cystolepiota mainly by structure of
the common veil. Fundamental character differences separating Lepiota from
the related Macrolepiota include fruitbody habitus, hymenophoral trama, spore
wall composition, and structure of the common veil. From similar Leucoagaricus
and Leucocoprinus the genus Lepiota differs in fruitbody habitus, spore wall
composition, presence of clamp-connections (most species) and structure of the
common veil. Lepiota is easily distinguished from Chlorophyllum by fruitbody
habitus, structure of the common veil, and whitish lamellae (for green-spored
Chlorophyllum species).
358 ... Kosakyan & al.
Section Lepiota
Lepiota clypeolaria (Bull.) P. Kumm., Fiihr. Pilzk.: 137. 1871. FIGURE 6
Descriptions and illustrations: Binyamini (1975: 147), Wasser (1980: 253-
254), Candusso & Lanzoni (1990: 168-172, pl. 15), Vellinga (2001: 116-117).
Hapitat: Solitary or in small groups, saprotrophic on the soil, in the fields and
broadleaf forests.
GENERAL DISTRIBUTION: Africa: Algeria, Morocco; Asia: Azerbaijan, China, Georgia,
Israel, Mongolia, Russia, Sri Lanka, Turkey; Australia; Europe: Austria, Belgium, British
Isles, Czech Republic, Denmark, Finland, France, Germany, Italy, Latvia, Lithuania,
Poland, Russia, Serbia, Spain, Ukraine; North America: Canada, Cuba, Mexico, USA;
South America: Chile.
MATERIAL EXAMINED: GH: Bar’am Wood, Quercus trees, 25.12.1972, leg. R. Neeman,
det. M. Moser, rev. S.P Wasser & M. Didukh (TELA-NM-72.464); SP: Tel Aviv University,
in grass, 19.04.1980, leg. & det. N. Binyamini, rev. M. Didukh, 20.03.2003 (TELA-NM-
80.645).
Notes: Lepiota clypeolaria is a polymorphic species. Modern species concepts
vary according to author. For instance, Israeli specimens and descriptions |
of species from Ukraine (Wasser 1980) appear to differ from those from
Netherlands (Vellinga 2001). Basic differences involve spore size, particularities
of the fruit body, odor, and taste.
Lepiota cortinarius J.E. Lange, Dansk. Bot. Ark. 2(3): 25. 1915. FIGURES 2, 6
PILEus 4-6 cm, conical when young, with inflexed margin, later applanate, with
broad umbo, at centre tomentose, around centre breaking up into squamules.
In young specimens squamules are brownish, later on around the center they
are paler, on cream background, immature fruit bodies with whitish velar
remnants at margin. LAMELLAE crowded, free from stipe, white to cream, with
whitish flocculose edge. STIPE 4-5 x 0.4-1.0 cm, with almost 2 cm wide bulb at
the base, hollow, whitish at apex, creamy-yellowish with few squamules below.
CONTEXT whitish, with strong flour smell. SPORE PRINT whitish.
BASIDIOSPORES 6.5-9.0 x 3.0-4.0 um, oblong ellipsoid, with lateral apiculus,
dextrinoid, not metachromatic in Cresyl Blue. Basip1A 4-spored, 15-30 x
5-8 um, clavate. PLEUROCYSTIDIA absent. CHEILOCYSTIDIA 20-50 x 6-15
um, mostly clavate or cylindrical-clavate. HYMENOPHORAL TRAMA regular.
PILEIPELLIS composed by long elements, tapering in apex, with brownish
pigment, mixed with basal short elements. STIPITIPELLIS composed by colorless
cylindrical hyphae. CLAMP-CONNECTIONS present in all tissues.
Hasirat: Solitary to gregarious, saprotrophic on soil, in grasslands.
GENERAL DISTRIBUTION: Asia: Israel; Europe: Denmark, France, Germany, Greece,
Lithuania, the Netherlands; North America: USA.
|
Lepiota species diversity in Israel ... 359
FIGURE 2. Lepiota cortinarius (HAI-525).
a — basidiomata, b — basidiospores, c — cheilocystidia. Bars = 10 um.
MATERIAL EXAMINED: CM: Mt. Carmel National Park, in grass, 08.12.1994, leg. & det.
S.P. Wasser (HAI-525).
Notes: Lepiota cortinarius, rare in Europe, is recorded here from Asia for the
first time.
Lepiota erminea (Fr.) P. Kumm., Fiihr. Pilzk.: 136. 1871. FIGURE 6
=Lepiota alba (Bres.) Sacc., Syll. fung. 5: 37. 1887.
Descriptions and illustrations: Binyamini (1984: 60, as Lepiota alba), Candusso
& Lanzoni (1990: 173-174, pl. 13a), Vellinga (2001: 118-119).
Hasirat: Solitary to gregarious, in small groups, saprotrophic on the soil, in
coniferous forests.
GENERAL DISTRIBUTION: Africa: Morocco; Asia: Georgia, Israel, Russia (Eastern Siberia),
Turkey; Europe: Belarus, British Isles, Czech Republic, Denmark, France, Germany,
Italy, Latvia, Lithuania, Poland, Spain.
360 ... Kosakyan & al.
MATERIAL EXAMINED: PP: Hadera, Iron wood, under Pinus sp. on the ground, 3.01.1978,
leg. & det. N. Binyamini, rev. M. Didukh (TELA-NM-78.414).
Notes: Lepiota erminea is rare in Israel. Two names exist in the literature
for a white species with fusiform spores: L. erminea and L. alba. The basic
differentiating criteria are the raphanoid smell and longer spores of L. erminea.
However, the latter trait is not very reliable, since L. alba spores vary significantly
(Babos 1979). Furthermore, the overlap of spore dimensions of the two species
— (12-)13-17(-20) x (5-)6-6.5(-7) um in L. alba vs. (13-)15-18(-21) x
5-7 um in L. erminea (Bon 1996) — can make separation of the two species
even more difficult, as also demonstrated in Huijsmanss (1943) and Vellinga's
(2001) studies of Dutch material. As a result, Vellinga (2001) regards smell and
spore size as insufficient to separate two species. The two species are currently
referred to L. erminea.
Lepiota ignivolvata Bousset & Joss. ex Joss., Riv. Micol. 33(1): 30. 1948. FIGURES 3, 6
Piteus 3-4.5 cm, with slightly umbonate brown centre, around centre with
brownish squamules on pale background. LAMELLAE free, crowded, white to
cream colored. STIPE 4-5 x 0.5-1.5 cm, with brown narrow ring, whitish cream,
discoloring orange when touched. CONTEXT white to creamy; smell strong, like
the rubber smell of L. cristata (Bolton) P. Kumm. SPorRE PRINT white.
BasIDIosPoREs 8.0-11(-13) x 5.0-6.5 um, amygdaliform, with fluorescent
content, dextrinoid, not metachromatic in Cresyl Blue. Basip1 4-spored, 20-
25(-30) x 6-8 um, clavate. PLEUROCYSTIDIA absent. CHEILOCYSTIDIA 15-35
x 8.0-15 um, clavate. PILEIPELLIS composed by long, brownish erect elements
70-300 um, mixed with rather short ones. CLAMP-CONNECTIONS present.
Hasirat: Solitary or in small groups, saprotrophic on soil, in grasslands.
GENERAL DISTRIBUTION: Asia: Israel; Europe: Denmark, France, Germany, Italy,
Luxemburg, Switzerland.
MATERIAL EXAMINED: CM: Mt. Carmel, Haifa University, National Park, in grass,
17.12.2000, leg. & det. S.P Wasser (HAI-SP-19).
Notes: Lepiota ignivolvata is recorded for the first time in Israel. It is also a new
species for Asian mycobiota. By habitus and smell it is similar to L. cristata.
It differs from the latter by distinct ring and shape of spores (truncate and
narrowly triangular in L. cristata). Not edible.
Lepiota oreadiformis Velen., Ceské Houby 1: 215. 1920. FIGURES 4, 6
Pireus 2-5 cm in diam., thin-fleshed, convex, plano-convex later on, with
umbo, yellowish-fulvous, skin-colored, darker in the center, paler towards the
margin, covered with slight flake like pruina in young fruit bodies, without
pruina later on; edge at first involute, straight and wavy later on. LAMELLAE
Lepiota species diversity in Israel ... 361
Chis chests
FiGurE 3. Lepiota ignivolvata (HAI-SP-19).
a — basidiospores, b - basidia, c - cheilocystidia, d - elements of the pileipellis. Bars = 10 um.
free, thin, white, later on with an ochroid tint. STIPE 2.5-—7 x 0.2-0.4 cm, central,
cylindrical, often slightly curved, fistulose in the center, whitish in the upper
part, concolorous with the pileus towards the base, covered with slightly flake
below the ring. RNG whitish, evanescent. CONTEXT white, getting pale-fulvous
tint below cuticle and in the base of the stipe. SPORE PRINT white.
BastpIospores 11.5-14 x 4.5-5.5 um, ellipsoid-amygdaloid, with lateral
apiculus, with one or some fluorescent drop or without it. BAsrp1a 4-spored,
35-45 x 10-11 um, cylindrical-clavate, clavate. PLEUROCYSTIDIA absent.
CHEILOCYSTIDIA 12-34 x 5-13 um, narrowly clavate, narrowly utriform,
some with a thickened wall. HYMENOPHORAL TRAMA regular. PILEIPELLIS
362 ... Kosakyan & al.
rn Ay RRR ij La cesesss
Dia SS ee x ©
coe
be
3
Figure 4. Lepiota oreadiformis (HAI-529).
a — basidiospores, b - basidia, c - cheilocystidia, d - elements of the pileipellis. Bars = 10um.
composed by erect elements, 50-150 x 5-12 um, sometimes mixed with short
clavate elements, often both type of elements are brown colored. STIPITIPELLIS
composed by 5-10 wm wide cylindrical yellowish elements. CLAMp-
CONNECTIONS present.
Hasirat: Solitary or in small groups, saprotrophic on soil, in grasslands.
GENERAL DISTRIBUTION: Asia: Azerbaijan, Israel, Russia (Eastern Siberia), Turkey;
Europe: British Isles, Estonia, France, Germany, Lithuania, Russia, the Netherlands.
MATERIAL EXAMINED: SP: Ramat Gan, in grassland, 06.02.2002, leg. & det. S.P Wasser &
M. Didukh. (HAI-529). FIGURE 5
Notes: Lepiota oreadiformis is recorded for the first time for Israel mycobiota.
It is close to L. pallida, which has septate pileipellis elements (Candusso
& Lanzoni 1990). Vellinga (2001) notes that a more yellowish coloration
and smaller sized spores distinguish dark specimens of L. erminea from
L. oreadiformis. Sometimes, L. oreadiformis may be confused with young
specimens of L. clypeolaria, which produces similar spores. L. clypeolaria can
be differentiated by a pileus cuticle that splits up in the distinct central disc
surrounded by discrete squamules in contrast to L. oreadiformis characterized
by an evenly colored pileal surface that uniformly splits up into small patches
(Vellinga 2001).
}
Lepiota species diversity in Israel ... 363
Lepiota pallida Locq. ex Bon & Candusso,
in Candusso & Lanzoni, Lepiota s.1. Fungi Europaei 4: 182.1990. | FiGuRES 5-6
PiLeus 0.8-1.9 cm in diam., very thin-fleshed (0.5-1.0 mm), hemispheric,
later plano-convex, with a small umbo, dark brown in the center, pale-brown,
whitish-ochre, brownish-red towards the margin, scaly-flaked, scales formed
by bundles of brownish and whitish hairs, coarse-fibrillose, dry with straight,
slightly fringed edge, often bearing velar remnants. LAMELLAE free, thin,
crowded, with an even edge, white, pale cream on drying. STIPE 2-4 x 0.2-0.3
cm, central, straight, sometimes slightly curved, thickened towards the base,
fistulose, whitish, smooth, covered with whitish-ochre pruina up to half of the
stipe. RING evanescent. CONTEXT thin, white, unchanging on autooxidation,
without a particular taste or odor. SPORE PRINT white.
BASIDIOSPORES 11.5-14.5 x 4.5-5.5 um, ellipsoid-amygdaloid, fusiform,
with lateral apiculus, with slight suprahilar depression, with or without one-
two fluorescent drops, smooth, thin-walled, dextrinoid, not metachromatic
in Cresyl Blue. Basrtp1a 4-spored, 24-32 x 6-7 um, clavate. PLEUROCYSTIDIA
absent. CHEILOCYSTIDIA 20-37 x 5-7 um, clavate or fusiform. HYMENOPHORAL
TRAMA regular. PILEIPELLIS consisting of obtuse or nearly cylindrical septate
hyphae 3-6 um in diam., long, not septate elements,150-250 x 5-10 um are
also present. CLAMP-CONNECTIONS present.
Hasitat: Mostly in small groups, saprotrophic on soils, in broadleaf forests.
GENERAL DISTRIBUTION: Africa: Morocco; Asia: Israel; Europe: France, Hungary, Italy,
Ukraine.
MATERIAL EXAMINED: PP: Hadera, under Eucalyptus, on the ground, in groups,
10.03.2002, leg. S.P. Wasser, det. M. Didukh (HAI-530).
Notes: Lepiota pallida is new for Israel and Asian mycobiota. This species, one
of the smallest Lepiota species, is rare throughout the world and previously
known from six localities in Europe and one in North Africa (Locquin 1945,
Kiihner & Romagnesi 1953, Babos 1969, Malencon & Bertault 1970, Wasser
1975). Our finding is the eighth report of this rare species. Wasser (1980) has
observed that L. pallida is found mostly in arid habitats.
In his original diagnosis, Locquin (1945) did not indicate cystidia; his name,
which was invalid due to the incomplete Latin description, was later validated
by Candusso & Lanzoni (1990). Although Hungarian specimens of L. pallida
collected by Babos (1969) also lack cystidia, Ukrainian (Wasser 1980), Italian
(Candusso & Lanzoni 1990), and Israeli specimens have cheilocystidia.
The habitat of the species differs depending on the country studied. In
Italy and France it grows under Cedrus and Robinia pseudoacacia (Candusso
& Lanzoni 1990), in Ukraine it grows in Pinus forest and under Robinia
pseudoacacia (Wasser 1980); in Israel, under Eucalyptus tree. Morphologically,
364 ... Kosakyan & al.
a
B.
eh
Rte
FiGure 5. Lepiota pallida (HAI-530).
a — basidiospores, b — basidia, c - cheilocystidia, d —- elements of the pileipellis. Bars = 10um.
the species resembles small representatives of Lepiota clypeolaria and
L. oreadiformis. From the latter species it differs mostly in septate pileus
covering elements.
Lepiota subgracilis Kiihner, Bull. Soc. mycol. France 52: 231. 1936. FIGURE 6
Description and illustration: Kosakyan et al. (2008: 66-68).
Hasirat: Solitary or in small groups, saprotrophic on the soil, in coniferous
-
forests.
GENERAL DISTRIBUTION: Asia: Israel, Russia (Far East-Primorye Territory), Sri Lanka,
Turkey; Europe: British Isles, Czech Republic, France, Hungary, Italy, the Netherlands,
Ukraine; North America: USA; South America: Argentina.
Lepiota species diversity in Israel ... 365
mi - L. clypeolaria
V -L. cortinarius
A -L. erminea
@ - L. ignivolvata
a L. oreadiformis
@ - 1. pallida
v - L. subgracilis
Figure 6. The distribution of representatives of section Lepiota in Israel.
MATERIAL EXAMINED: Israel. CM: Ramat Hanadiv, in Pinus stand, leg. Y. Ur, 19.02.2007,
det. A. Kosakyan (HAI 328).
Notes: Lepiota subgracilis is a relatively common species. For Israel it is recorded
for first time (Kosakyan et al. 2008). L. subgracilis is a quite variable species. Our
specimens differ from those described in Wasser (1980) in producing smaller
pilei and larger, more variable cystidia. Candusso & Lanzoni (1990) describe
specimens with smaller spores, while Vellinga (2001) gives larger pileus, stipe,
and spore sizes and shorter, narrower pileipellis hyphae for L. subgracilis.
Section Ovisporae (J.E. Lange) Kuhner
Lepiota brunneoincarnata Chodat & C. Martin,
Bull. Soc. Bot. Geneve, 2 sér. 5: 222. (1889). FIGURE 11
Descriptions and illustrations: Binyamini (1975: 149), Wasser (1980: 248-
252), Candusso & Lanzoni (1990: 248-252), Vellinga (2001: 127).
Hasirat: Gregarious or more rarely solitary, saprotrophic on soil, in parks,
grasslands, coniferous and broadleaf forests.
GENERAL DISTRIBUTION: Africa: Algeria, Morocco; Asia: Azerbaijan, India, Israel;
Europe: Austria, British Isles, Czech Republic, Denmark, Estonia, France, Germany,
Poland, Sweden, Switzerland, the Netherlands, Ukraine.
366 ... Kosakyan & al.
DISTRIBUTION IN ISRAEL: SP: Hadera, under Eucalyptus trees, 14.12.1972, leg. & det.
N. Binyamini (TELA-NM 72.348); Tel Aviv, Hayargon Park, under Cupressus tree,
25.02.1973, leg. & det. N. Binyamini (TELA-N. 73.168); Ilanot, on lawn, 28.09.1978,
leg. & det. N. Binyamini (TELA-N. 78.348); Tel Aviv, Rosh-Ha‘ayin Park, on lawn,
under Eucalyptus trees, 18.12.1989, leg. N. Binyamini and A. Prusbul, det. N. Binyamini
(TELA-N. 89.276).
MATERIAL EXAMINED: CM: Mt. Carmel, Haifa University National Park, on grass,
19.12.2000, leg. S. Tovbin, det. S.P. Wasser (HAI-SP-517); Mt. Carmel, Nahal Nesher,
park, under Casuarina sp. 12.04.2003, leg. S.P. Wasser, det. M. Didukh (HAI- 524).
Notes: Lepiota brunneoincarnata is a very toxic species. It is known to contain
high level of amatoxins. It contains a- and y-amanitins — up to 3.5mg/g; a lethal
dose for humans is approximately 50 to 300g of fresh mushroom (Meunier et
ala 995).
Lepiota brunneolilacea Bon & Boiffard,
Bull. trimest. Soc. mycol. France 88: 18 (1972). FIGURE 11
Descriptions and illustrations: Candusso & Lanzoni (1990: 252-255),
Courtecuisse & Duhem (1994: 224, fig. 687), Didukh et al. (2002: 44).
HasirTat: Solitary or in small groups, saprotrophic on soil, in parks, in Pinus
groves.
GENERAL DISTRIBUTION: Asia: Israel; Europe: France, Italy, Spain, the Netherlands,
Ukraine; North America: Canada.
MATERIAL EXAMINED: CM: Mt. Carmel National Park, Haifa University, under Pinus
halepensis, 20.12.2001, leg. K. Tzukor, det. M. Didukh (HAI SP-17).
Notes: L. brunneolilacea was described by Bon & Boiffard (1972) from
the Atlantic and Mediterranean coasts of France, growing on dunes amid
Ammophila arenaria. This species is new for Asian mycobiota.
Lepiota echinella Quél. & G.E. Bernard, Bull. Soc. mycol. France 4: pl. 1, fig. 2
(1888). FIGURES 7, 11
PiLeus 1.5 cm in diam., thin-fleshed, hemispheric, later on convex-plane, with
a small umbo, brownish-reddish on white background, covered with uplifted
scales, thin, edge involute, wavy. LAMELLAE free, thin, crowded, white, with
an uneven eroded concolorous edge. STIPE 2-4 x 0.1-0.2 cm, central, pinkish,
covered with darker colored fibrils, brownish at the base, with floccose-scaly
pruina. RING brownish, disappears at early stages, leaving a nearly indiscernible
zone. CONTEXT white, slightly browning on autooxidation, without particular
odor. SPORE PRINT whitish.
BASIDIOSPORES 5-6.5 x 3.5-4 um, ellipsoid, with fluorescent content, weakly
dextrinoid, not metachromatic in Cresyl Blue. BAsrp1a 4-spored, 20-23 x 6-7
tum, clavate. PLEUROCYSTIDIA absent. CHEILOCYSTIDIA vary in shape: fusiform,
Lepiota species diversity in Israel ... 367
FiGure 7. Lepiota echinella (HAI-M-528).
a — basidiospores, b — basidia , c — basidia, d — cheilocystidia, e - elements of the pileipellis.
Bars = 10um.
lageniform, utriform, thin-walled, hyaline, 20-25 x 6-8 um. HYMENOPHORAL
TRAMA regular. PILEIPELLIS composed of erect, long, cylindrical brownish
elements and short, narrowly clavate to cylindrical elements 15-50 x 5-12
lum. STIPITIPELLIS composed of narrow cylindrical hyphae, sometimes with a
brown pigment. CLAMP-CONNECTIONS present in all tissues.
HasirTat: Gregarious in small groups, saprotrophic on soils, in grasslands.
GENERAL DISTRIBUTION: Africa: Morocco; Asia: Israel; Europe: British Isles, Czech
Republic, Denmark, France, Germany, Hungary, Italy, Poland, the Netherlands,
Ukraine.
MATERIAL EXAMINED: SP: Ramat-Gan, park, on the lawns, 28.03.2003, leg. Y. Ur, det.
M. Didukh (HAI-M-528).
368 ... Kosakyan & al.
Notes: Lepiota echinella is a rare species in European countries. It is recorded
for the first time in Israel. Vellinga (2001) notes that L. lepida Guinb. & M.
Bodin comes very close to L. echinella, with its spores being slightly bigger
and cheilocystidia more variable in size and shape in L. echinella. L. locquinii
is likewise similar but can be differentiated by its smaller pileus and differently
shaped cheilocystidia (clavate in L. locquinii and lageniform or utriform in
L. echinella).
Lepiota forquignonii Quél., |
Compt. Rend. Assoc. Frang. Avance. Sci. 13: 277. (1885). FIGURES 8, 11
PiLeus 1-4 cm in diam., thin-fleshed, hemispherical, campanulate when young,
later applanate with an obtuse umbo, dark grey, smooth in the center; apart
from the central calotte the cuticle is broken up into variously shaped, grayish,
brownish, adpressed, concentric squamules on the white background, radially
fibrillose towards the margin; edge involute, thin. LAMELLAE free, ventricose,
not very crowded, thick, with white eroded edge. STIPE 2.5-5 x 0.4-0.6 cm,
cylindrical or tapering towards the apex, slightly widening towards the base, |
whitish, pale grayish-greenish, without a ring, covered with pale fulvous fibrils
in the lower part, broadly fistulous. CONTEXT cotton-like, whitish, pale pink in
stipe and pileus. Odor unpleasant, as in L. cristata. SPORE PRINT whitish.
BASIDIOSPORES 6-8 x 3.0-5 ym, oval, slightly dextrinoid, not metachromatic
in Cresyl Blue. Basrp1a 4-spored, 14-20 x 6-8 um, clavate, two-spored
basidia can occur closer to the edge of the gills. PLEUROCysTIDIA absent.
CHEILOCYSTIDIA numerous, collected in bundles, variously shaped, 20-32 x
4.5-8.0 um, nearly cylindrical, narrowly fusiform, narrowly clavate, some are
capitate; with clamp-connections at the base. HYMENOPHORAL TRAMA regular.
PILEIPELLIS composed by long erect elements 80-400 x 5-10 um in diam.,
collected in bundles, mostly thick-walled, with yellowish parietal pigment,
tapering towards the apex, with slightly fusiform base, with clamps at the base
of some elements; few narrowly clavate elements 24-28 x 7.2-10 um occur
among long elements. STIPITIPELLIS composed by narrow cylindrical hyphae
3.0-5.0 um wide. CLAMP-CONNECTIONS present in all tissue.
Hasirat: In small groups, saprotrophic on soil, in broadleaf forests.
GENERAL DISTRIBUTION: Africa: Morocco; Asia: Israel, Turkey; Europe: Italy, Sweden,
the Netherlands; North America: USA; South America: Brazil.
MATERIAL EXAMINED: SP: Hadera, park, under Eucalyptus sp., on the ground, 09.12.2004,
leg. Y. Ur, det. M. Didukh (HAI-520).
Notes: Lepiota forquignonii is new species for Israel. It is uncommon in
Europe, where it is known primarily from the temperate regions and only rarely
found in the warmer regionss (Vellinga 2001). In view of this, the finding of
Lepiota species diversity in Israel ... 369
Figure 8. Lepiota forquignonii (HAI-520).
a — basidiospores, b — basidia, c — basidia, d - cheilocystidia, e - elements of the pileipellis.
Bars = 10um.
L. forquignonii in warmer Israeli climate contributes to the knowledge about
adaptive abilities of the species.
Lepiota helveola Bres., Fung. trident. 1(1): 15. 1882 (1881). FIGURES 9, 11
Pieus 1.5-4cm in diam., thin-fleshed, hemispheric, plano-convex or applanate
later on, with a broad umbo, dirty grayish-pinkish, grayish-reddish, reddish-
ochraceous-brownish, slightly fibrillose, covered with adpressed tiny dark
squamules, between which white flesh can be seen; edge thin, involute, later on
straight, wavy, often cracked. LAMELLAE free, thin, white, with a yellowish tint
later on. STIPE 2-4(5) x 0.3-0.8 cm, central, cylindrical, fistulose, concolorous
with the pileus or paler, smooth above the ring, with flake-like pruina below
370 ... Kosakyan & al.
— ght
FiGuRE 9. Lepiota helveola (HAI-531).
a — basidiospores, b — basidia, c — basidia, d — cheilocystidia, e - elements of the pileipellis.
Bars = 15um.
the ring. RNG upper, funnel-shaped, whitish on the upper side, with lower
side concolorous with the pileus. CONTEXT white, peripheral layer turns red on
autooxidation, without a particular taste or odor. SPORE PRINT white.
BASIDIOSPORES 7-8 x 3.5-4 um, colorless, ellipsoid, with lateral apiculus
and fluorescent contents. Bastp1a 4-spored, 25-32 x 8-10 um, clavate.
PLEUROCYSTIDIA absent. CHEILOCYSTIDIA 28-34 x 8-11 um, narrowly
clavate. HYMENOPHORAL TRAMA regular. PILEIPELLIS composed by clavate
long elements 50-100 x 10-20 um, with pigmentation in walls and at the base, |
mixed with rather short elements. CLAMP-CONNECTIONS present. |
Hapirtat: Solitary or in small groups in open areas, in coniferous forests. |
GENERAL DISTRIBUTION: Africa: Algeria, Morocco; Asia: Azerbaijan, Israel, Japan;
Europe: Czech Republic, France, Italy, Portugal, Spain, Ukraine.
Lepiota species diversity in Israel ... 371
MATERIAL EXAMINED: GH: Banias, under Pinus halepensis, 26.01.2003, leg. & det. S.P.
Wasser & M. Didukh (HAI-531).
Notes: Lepiota helveola is a very rare species in Israel. Binyamini (1975), who
first reported L. helveola for the country, did not provide information on the
collection site or where the dried specimens were deposited. Poisonous.
C— qv
FiGurE 10. Lepiota locquinii (HAI-518).
a — basidiospores, b — basidia, c — basidia, d - cheilocystidia, e - elements of the pileipellis.
Bars = 10um.
Lepiota locquinii Bon, Doc. Mycol. 16(61): 20. 1985. FIGURES 10-11
PiLgus 1-1.5 cm in diam., thin-fleshed, hemispheric, with a small dark brown
umbo, covered with scales formed upon cuticle breaking; with involuted,
372 ... Kosakyan & al.
wavy. LAMELLAE free, thin, crowded, white-cream, with an uneven eroded
concolorous edge. STIPE 1.5-2 x 0.1-0.2 cm, central, creamish-pinkish, covered
with few fibrils, brownish-reddish at the base. RING disappears at early stages.
CONTEXT white, without any particular odor. SPORE PRINT whitish.
BASIDIOSPORES 7.5-9 x 4.5-5 um, colorless, ellipsoid, with fluorescent context,
weakly dextrinoid, not metachromatic in Cresyl Blue. BAsip1a 4-spored, 20-23
x 6-7 um, clavate. PLEUROCYSTIDIA absent. CHEILOCYSTIDIA 18-25 x 5-8
um, mostly clavate. HYMENOPHORAL TRAMA regular. PILEIPELLIS composed
by erect, long, cylindrical elements 50-100(150) x 8-10, with some narrowly
clavate to cylindrical elements at the base. STIPITIPELLIS composed by narrow
cylindrical hyphae, sometimes with brown pigment. CLAMP-CONNECTIONS
present in all tissues.
HaBITAT: Gregarious, in broadleaf and coniferous forests, on humus-rich
soils.
GENERAL DISTRIBUTION: Europe: Italy, France; Asia: Israel.
MATERIAL EXAMINED: CM: Mt. Carmel, near Nahal Oren, in mixed wood of Pinus and
Quercus, 25.12.1996, leg. & det. S. P. Wasser (HAI-518).
Note: Lepiota locquinii is very rare worldwide and recorded here for the
first time from Israel and Asia. L. locquinii is very similar to L. echinella,
from which it can be differentiated by its smaller, paler pileus and differently
shaped cheilocystidia (clavate in L. locquinii and lageniform to utriform in
L. echinella).
Lepiota subincarnata J.E. Lange, Fl. Agaric. Danic. 5(Append.): v. 1940. FiGURE 11
Descriptions and illustrations: Binyamini (1976: 70), Candusso & Lanzoni
(1990: 244-247, pl. 25a), Vellinga (2001: 127).
HasitTatT: Gregarious, in small groups, saprotrophic on soils, in city-parks,
gardens, in broadleaf forests.
GENERAL DISTRIBUTION: Asia: Israel, Turkey, Europe: France, Germany, Italy, Poland;
North America: USA.
MATERIAL EXAMINED: SP: Hadera, Iron wood, mixed forest of Quercus and Pinus trees,
18.12.1973, leg. & det. N. Binyamini, rev. M. Didukh & S.P Wasser (TELA-N. 73.293);
Tel Aviv, Hayargon Park, on lawn, 26.02.1974, leg. & det. N. Binyamini, rev. M. Didukh
& S.P. Wasser (TELA-N. 74.435); Miqwe-Yisrael, under Eucalyptus sp., leg. & det. N.
Binyamini, 10.12.1974, rev. M. Didukh & S.P. Wasser (TELA-N. 16.182); Hadera Park,
under Eucalyptus, on the ground, 09.01.2004, leg. Y. Ur, det. M. Didukh (HAI-521); CM:
Mt. Carmel, Carmel National Park, on grass, date 15.01.2003, leg. & det. S.P. Wasser
(HAI- SP-23).
Notes: Lepiota subincarnata is not common in Asia, but has been found in
Israel several times. This very toxic species contains amanitins (Bresinsky &
Besl 1990).
a ae
Lepiota species diversity in Israel ... 373
Vv - L. brunneoincarnata
| Jf - L. brunneolilacea
@ - L. echinella
we - L. forquignonii
| w - L. helveola
A - L. locquinii
| > - L. subincarnata
FiGureE 11. The distribution of representatives of section Ovisporae in Israel.
Section Stenosporae (J.E. Lange) Kuhner
Lepiota castanea Quél.,
Compt. Rend. Assoc. Frang. Avance. Sci. 9: 661. 1881. FIGURE 12
Descriptions and illustrations: Binyamini (1984: 60), Candusso & Lanzoni
(1990: 198-202), Vellinga (2001: 130-131).
Hasirat: Solitary to gregarious, saprotrophic on soil, in coniferous forests.
GENERAL DISTRIBUTION: Africa: Morocco; Asia: Armenia, Azerbaijan, Israel, Japan,
Russia, Turkey; Europe: British Isles, Czech Republic, Denmark, France, Germany, Italy,
Latvia, Lithuania, Poland, Russia, the Netherlands; New Guinea.
MATERIAL EXAMINED: PP: Hadera, under Pinus sp., Wadi Ara, Iron Wood, 16.01.1979,
leg. student, det. N. Binyamini, rev. S.P. Wasser & M. Didukh (TELA-N. 79.188).
Notes: Lepiota castanea, which is a rare in Israel, possesses medical properties.
L. castanea exhibits antitumor activity against Sarcoma 180 and Ehrlich
carcinoma at different inhibition rates (Ohta et al. 1998).
374 ... Kosakyan & al.
@ - L. castanea
FiGuRE 12. The distribution of representatives of section Stenosporae in Israel.
Section Cristatae Kuhner ex Wasser
Lepiota apatelia Vellinga & Huijser, Belg. J. Bot. 131: 196. 1999 (‘1998’). FIGURE 13
Description and illustration: Kosakyan et al.(2008: 59-74).
Hapsitat: Gregarious, often in big groups, saprotrophic on soil, in broadleaf
forests.
GENERAL DISTRIBUTION: Asia: Israel; Europe: Austria, Germany, Italy; the Netherlands.
MATERIAL EXAMINED: UG: Safsufa forest, under Quercus calliprinos, leg. Y. Ur,
19.11.2006, det. A. Kosakyan (HAI 33).
Notes: L. apatelia is a very rare species that was described recently from the
Netherlands by Vellinga & Huijser (1999). It is new species for Israel and for
Asian mycobiota (Kosakyan et al. 2008). L. apatelia is macroscopically very
similar to L. cristata, from which it is easily distinguished by basidiospore
shape, absence of distinct annulus, and different smell. It is also very close
to L. cristatoides Einhell., from which it differs by uneven and orange-brown
pileus, absence of annulus, fruitbody size, and spore microchemical reactivity
(Vellinga & Huijser 1999; Vellinga 2001). The similar Lepiota thiersii Sundb.,
described from California (Sundberg 1989), can be differentiated by spores that
Lepiota species diversity in Israel ... 375
| & - L. apatelia
| @ -L. lilacea
me - L. micropholis
FiGuRE 13. The distribution of representatives of section Cristatae in Israel.
are not dextrinoid, a less well-developed partial veil, possession of clavate to
sphaeropedunculate pileipellis elements, and a farinaceous odor (Vellinga &
Huijser 1999).
Lepiota lilacea Bres., Fung. trident. 2(8-10): 3. 1892. FIGURE 13
Descriptions and illustrations: Binyamini (1975: 150), Wasser (1980: 246),
Candusso & Lanzoni (1990: 308-311), Vellinga (2001: 140-141).
Hasitat: Solitary or in small groups, saprotrophic on soils, often in parks, also
in coniferous forests.
GENERAL DISTRIBUTION: Africa: Algeria, Ghana, Morocco, Zaire; Asia: Azerbaijan,
China, Iran, Israel; Europe: British Isles, Czech Republic, Denmark, France, Germany,
Italy, Poland, Serbia, the Netherlands, Ukraine; South America: Venezuela.
MATERIAL EXAMINED: SP: Tel Aviv, Hayarqon Park, on lawn, 17.10.1972, leg. & det. N.
Binyamini, rev. M. Didukh & S.P. Wasser (TELA-N. 72.220); Iron Wood, under Pinus,
24.02.1973, leg. & det. N. Binyamini, rev. M. Didukh & S.P. Wasser (TELA-N. 73.167).
Notes: Lepiota lilacea is a very rare species for Israel. Vellinga (2001) notes that
it can be very variable, especially with respect to basidiocarp size and color.
Dark variants may resemble L. felina (Pers.) P. Karst in annular structure and
376 ... Kosakyan & al.
the discrete disc and pileus cuticle squamules, but the two species differ in
pileipellis structure and spore size (Vellinga 2001).
Lepiota micropholis (Berk. & Broome) Sacc., Syll. fung. 5: 61. 1887. FIGURE 13
Descriptions and illustrations: (Binyamini 1975: 150), Saccardo (1887: 61),
Pegler (1972: 160, fig. 1/4 a—d).
Hapsitat: Solitary or in small groups, saprotrophic on soil, often in parks, also
in broadleaf forests.
GENERAL DISTRIBUTION: Africa: Kenya, Morocco; Asia: Japan, India, Israel, South
Korea, Sri Lanka; Europe: Hungary, Ukraine; South America: Brazil, Venezuela.
MATERIAL EXAMINED: SP: Ramat Hasharon, in orchard near orange tree, 07.12.1974,
leg. & det. N. Binyamini, rev. S.P. Wasser & M. Didukh (TELA-N. 74.457); Ramat-Gan,
in Eucalyptus groves, 24.01.1978, leg. & det. N. Binyamini, rev. S.P. Wasser & M. Didukh
(TELA-N. 79.270).
Notes: Lepiota micropholis is a rare species for Israeli eae tee previously
known from only two Israeli reports.
Acknowledgments
We thank the Dean of Graduate Studies Authority of Haifa University, Professor
Sophia Menache, for supporting our research and this publication. Also, we thank Dr.
I. Zmitrovich (Russia), Dr. E. Vellinga (USA), Dr. P. Volz (USA), Dr. S. Pennycook (New
Zealand), and Dr. L. Norvell (USA) for critical reviews of the manuscript.
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MY COTAXON
Volume 105, pp. 379-386 July-September 2008
Studies in lichens and lichenicolous fungi:
notes on some taxa from North America
JAMES C. LENDEMER’ , JANA KOCOURKOVA? & KERRY KNUDSEN?
jlendemer@nybg.org
*Cryptogamic Herbarium, Institute of Systematic Botany
The New York Botanical Garden, Bronx, NY, 10458-5126, USA
*National Museum, Dept. of Mycology
Vaclavské nam. 68, 115 79 Praha, Czech Republic
>The Herbarium, Dept. of Botany and Plant Sciences
University of California, Riverside, CA, 92521-0124, USA
Abstract — Acarospora instrata is placed in synonymy with A. obpallens. Cyphelium
brachysporum is removed from synonymy with C. notarisii and considered endemic
to the chaparral belt of southern California, USA. Amandinea insperata, Cercidospora
cladoniicola, Chaenothecopsis kalbii, and Usnea diffracta are reported for the first time
from North America.
Key words — Appalachian Mountains, coastal plain, fire frequency, New Mexico,
tropical disjuncts
1. Acarospora obpallens (Nyl. ex Hasse) Zahlbr., Beihefte. Bot. Centralbl., 13: 161.
1902.
Syn. nov. Acarospora instrata H. Magn., Kongl. Svenska Vetensk. HandL., ser. 3, 7(4):
279. 1929. TYPE: U.S.A., New Mexico, C.J. Harnk s.n. (W!, holotype).
Recently, the holotype of Acarospora instrata was examined in the context of
continuing revisionary work on Acarospora in North America. The holotype is a
typical faveolate, epruinose specimen of A. obpallens on silicate-rich rock which
is common in southern California and the mountains of the Sonoran Desert in
Arizona (Knudsen 2008). The synonymy is made here. Acarospora obpallens
is now documented from the New Mexican lichen biota. For descriptions and
pictures of A. obpallens see Knudsen (2008) and Knudsen et al. (2008).
2. Amandinea insperata (Nyl.) H. Mayrhofer & Ropin, in Mayrhofer et al.,
Muelleria, 12: 190. 2000 (“1999”).
During routine fieldwork in Nantahala National Forest in the southern
Appalachian Mountains of eastern North America, the first author collected an
380 ... Lendemer, Kocourkova & Knudsen
odd Amandinea that did not seem to represent any species presently reported
from the region. In light of the recent discovery of Buellia japonica (Tuck.) Tuck.,
a sub-tropical species, in a rich cove forest in adjacent Gorges State Park (Sheard
et al. 2008) a search of the additional tropical species treated by Marbach (2000)
was made. The material perfectly fits the description of Amandinea insperata,
a species with a pan-tropical distribution similar to that of B. japonica. The
species is presently known from South America (Argentina, Brazil, Ecuador,
Columbia, Paraguay, and Uruguay), Central America (Guatemala), Africa
(Kenya, Macronesia), and Australia. The species is similar to B. japonica in
that it is most likely to be confused with a species of Rinodina because of its
ascospores with elongated angular lumina (i.e. not Buellia-type). The brown
hypothecium, black-capped paraphyses, and filiform conidia clearly distinguish
the species from any Rinodina species in the region. This is the first report of
the species from North America.
SPECIMEN EXAMINED. — U.S.A. NORTH CAROLINA. JACKSON CO.: Nantahala
National Forest, Chattooga Wild and Scenic River / Ellicott Rock Wilderness, above
Fowler Creek, just S of Bull Pen Road, ~4.5 mi SE of Highlands, elev. 3000 ft., overgrown
homestead site along stream with black locust (Robinia pseudoacacia), apple (Malus),
and alder (Alnus) and large gneiss boulders, on branches of black locust, 18.ix.2006, J.C.
Lendemer et al. 7891 (NY).
3. Cercidospora cladoniicola Alstrup, Graphis Scripta 8: 26. 1997. TYPE: Norway,
Nord-Trondelag, Flatanger, Roythaugfjellet, UTM 32W NS 8548-49, map 1623 I,
alt. 40-120 m, ravine with vertical rock and coastal deciduous forest, on Cladonia
arbuscula, 27.vii.1993, V. Alstrup (C, holotype).
In the material collected by the second and third authors during a field study of
Santa Rosa Island in southern California in July 2007, we found the lichenicolous
fungus Cercidospora cladoniicola totally bleaching the host squamules of
Cladonia pyxidata (L.) Hoffm. The fungus is a parasite causing severe damage
to the host thalli including deformation of the podetia. It is known from various
Cladonia species, mostly Cladonia sect. Cladina such as C. arbuscula (Wallr.)
Flot., C. mitis Sandst., and C. portentosa (Dufour) Coem., but it has also been
reported on C. pocillum (Ach.) O.J. Rich. and C. symphycarpia (Flérke) Fr.
It is a widely distributed but rare species in the Northern Hemisphere,
previously reported from Norway (Alstrup 1997, type locality), Denmark
(Alstrup & Svane 1998), British Isles (Coppins 1998), Russia (Zhurbenko
2004) and Svalbard (Zhurbenko & Alstrup 2004). We report it new to North
America. ;
In the genus, Cercidospora cladoniicola belongs to the group of species with 8-
spored asciand 3-septate ascospores together with C. lecidomae Zhurb. & Triebel
and C. thamnoliicola thlen (Ihlen & Wedin 2007). Cercidospora cladoniicola
differs from both species in having ascospores brownish when mature and in
Lichens & lichenicolous fungi (North America) ... 381
its host. Moreover, from C. lecidomae, C. cladoniicola differs in having brown
peridium versus emerald to glaucous green peridium and in having regularly
3-septate instead of 1-3 septate ascospores, which are not narrow to attenuated
in lower part (Zhurbenko & Triebel 2003), and from C. thamnoliicola in larger
ascospores (13—)16-—18-20(-21) x (3—)4—4.5-5um versus 11-14(-16) x 4-6um
(Zhurbenko & Alstrup 2004). Other 3-septate Cercidospora species, C. soror
Obermayer & Triebel (Hafellner & Obermayer 1995) and C. stereocaulorum
(Arnold) Hafellner (Hafellner 1987), have 4-spored asci.
SPECIMEN EXAMINED. - U.S.A. CALIFORNIA. SANTA BARBARA CO.: Channel
Islands, north slope of Black Mountain, first canyon west of truck trail, alt. 267 m.,
33°58'58"N 120°4’4°W, senescent Bishop pine woods and chaparral, on Cladonia
pyxidata on soil, 18.vii.2007, J. Kocourkova & K. Knudsen s.n. (PRM 909679).
4. Chaenothecopsis kalbii Tibell & K. Ryman, Nova Hedwigia, 60: 206. 1995. TYPE:
Brazil, Mato Grosso do Sul, 50 km SW of Campo Grande, 1979, K. Kalb s.n. (hb.
Kalb, holotype).
Chaenothecopsis kalbii is a lichenicolous fungus occurring on the thallus of
Lecanora caesiorubella Ach. (Tibell 1996). The species was originally described
from Brazil, and is also known from Baja California Sur, Mexico and tropical
Australia (Tibell 1996). The discovery of this species in the coastal plain of the
southeastern United States is not unexpected, particularly in light of the number
of species with similar sub-tropical or pan-tropical distributions that have been
recently reported from that region (Beeching 2007, Lendemer 2008, Lendemer
& Knudsen 2008, Lendemer & Lumbsch in press, Lendemer & Yahr 2004).
Because of its short, black ascomata with a pale stalk, 1-septate ascospores
(9-11 x 4-5.5 um), and its lichenicolous habit (on otherwise pale thalli of
Lecanora caesiorubella) C. kalbii is a distinctive species easily recognized in
the field. Most likely, searching more host thalli will lead to the discovery of
additional populations in North America.
SPECIMEN EXAMINED. - U.S.A. NORTH CAROLINA. CARTERET CO.: Cape
Lookout National Seashore, Shackleford Banks, maritime forest, on thallus of Lecanora
caesiorubella, 19.iii.2003, R.C. Harris 47195 (NY).
5. Cyphelium brachysporum Nadvy., Ann. Myc., 40: 133. 1942. TYPE: US.A.,
California, Riverside Co., near Murietta, 1903, H.E. Hasse s.n. (PRM!, holotype;
BRA isotype).
In 1903 Herman Hasse collected a yellow Cyphelium on the wood of a dead
shrub in southern California in western Riverside County near Murrieta. Based
on the concepts of the period, he identified the specimen as Cyphelium tigillare
(Ach.) Ach. In 1905 he returned to western Riverside County and according
to labels collected in Murrieta more specimens of this Cyphelium on the dead
wood of the common chaparral shrub Adenostoma fasciculatum Hook. & Arn.
382 ... Lendemer, Kocourkova & Knudsen
In his flora he reported it as occurring “on dead shrubs near Murrieta” (Hasse
1913). In this. general area in the Temecula Valley and in the surrounding
foothills, chaparral containing Adenostoma fasciculatum occurs at elevations
from approximately 300 to 700 meters.
One of Hasse’s 1903 Murrieta collections ended up in a private herbarium in
Czechoslovakia. During World War II, the lichenologist Nadvornik (1942) saw
this specimen and realized it was new to science based on the small size of its
one-septate ascospores and described it as Cyphelium brachysporum.
Studying a single isotype of the 1903 collection, Tibell synonymized
Cyphelium brachysporum with C. notarisii (Tul.) Blomb. & Forssell (Tibell et
al. 2003). He based this synonymy on the speculation that the ascospores of
C. brachysporum described by Nadvornik were immature, and that if they were
mature that would fall within the range of the ascospores of C. notarisii (which
are submuriform/muriform).
Based on our study of the holotype, three later collections from 1905 by
Hasse, and one modern collection we reject the synonymy of Cyphelium
brachysporum with C. notarisii. Cyphelium notarisii is a well delineated taxon,
easily recognized by its large, smooth, muriform ascospores (17-24 x 13-18 ©
uum). The protologue of C. brachysporum reports 1-septate ascospores (11-13
x 8.5-10 um). Our study of five specimens, including the holotype, revealed
broadly ellipsoid ascospores ({13.5] -14.1- [14.7] x [10.0] -10.6-[11.3]um;
n=40, average +/— 1.96SE) overall slightly larger than the measurements given
in the protologue, but still outside of the range of C. notarisii, and with a rare
additional longitudinal division in the upper cell creating a sub-muriform
ascospore. The ascospores of C. brachysporum are also minutely punctate and
not constricted at the septum and the thallus contains rhizocarpic acid. The
infrequent formation of an additional longitudinal septum in the upper cell of
the ascospores led Tibell to assume they were immature muriform ascospores.
While we observed this character in one collection from 1905 (NY), this was
neither observed in the holotype nor the two additional collections from 1905
(FH) or the modern collection. It is our opinion that the mature ascospores
in C. brachysporum are 1-septate, only occasionally becoming submuriform.
The few submuriform ascospores we observed in the specimen at NY were
still smaller in size than those of C. notarisii. Further, we are unaware of any
collections of C. notarisii from southern California other than those we here
refer to C. brachysporum. Otherwise, C. brachysporum is similar in appearance
to C. tigillare with apothecia immersed in thalline warts, the exciple thin
throughout, rather than sessile like C. pinicola Tibell with the exciple strongly
thickened at the base.
The rarity of Cyphelium brachysporum is explained by its occurrence on
the unburned wood of dead chaparral shrubs at low elevations. For instance,
before widespread urbanization and urban sprawl in southern California, old-
Lichens & lichenicolous fungi (North America) ... 383
growth chaparral was common. In old-growth chaparral the shrubs at ages
of thirty-to-sixty years become senescent and die. It is on the wood of these
dead shrubs that C. brachysporum was first collected. In the last hundred years
anthropogenic fires have become common in the southern California chaparral
with fire frequencies as low as every twenty years. Old-growth chaparral is now
rare (Knudsen 2006). The single modern collection of C. brachysporum was
made in old-growth chaparral in San Luis Obispo County in Los Osos on the
dead wood of Morro Manzanita, Arctostaphylos morroensis Wiesl. & B. Schreib.
This is a unique chaparral type, Morro Bay chaparral, restricted to the local
sand dunes below 200 meters, that has very low natural fire frequency. This
dune chaparral also supports the only known populations of Sulcaria isidiifera
Brodo.
ADDITIONAL SPECIMENS EXAMINED. -— U.S.A. CALIFORNIA. RIVERSIDE CO.:
Murietta, 1905, H.E. Hasse s.n. (FH [2 specimens], NY); SAN LUIS OBISPO CO.: Los
Osos, Los Osos Oaks State Reserve, 57 m, 16.xii. 2005, Knudsen et. al 4615 (UCR).
6. Leptogium juressianum Tav., Portug. Acta Biol. Ser. B, 3(1-2): 68. 1950.
Recently, while determining specimens from the southern Appalachian
Mountains collected by Sean Beeching the first author encountered an unusual
isidiate Leptogium specimen with an arachnoid tomentum on both the upper
and lower surfaces of the thallus. As this type of tomentum is unusual in the
genus Leptogium and it rarely occurs on the upper surface of the thallus as
well as the lower surface, a search of the relevant literature (Jorgensen & James
1983) quickly led to the name L. juressianum. Since according to Jorgensen and
James (1983) L. juressianum is a rare species known only from hyperoceanic
areas of Europe (Portugal and Ireland) a duplicate was sent to P.M. Jorgensen
who kindly confirmed the identification. Leptogium juressianum is thus here
reported for the first time form North America.
A subsequent visit by the first author, Sean Beeching, and Malcolm Hodges
to the locality where L. juressianum was found yielded no further material of
the species. The locality where L. juressianum was found is a middle-elevation
serpentine barren, a habitat that is rare in the southern Appalachian Mountains
and has previously yielded other rare and endemic species (see Lendemer 2007
for a discussion).
SPECIMEN EXAMINED. — U.S.A. NORTH CAROLINA. CLAY CO.: Nantahala National
Forest, vicinity of Buck Creek Barrens, 2006, S.Q. Beeching s.n. (NY).
7. Usnea diffracta Vain., Bot. Mag. Tokyo, 35: 45. 1921. TYPE: Japan, Hokkaido,
Prov. Kushiro, on Betula, 19.viii.1918, A. Yasuda 231 (TUR, holotype).
While organizing the oversized specimens of lichens in the herbarium of the
Academy of Natural Sciences of Philadelphia (PH), the first author came across
an old folder of Usnea specimens that had apparently not been seen by any
384 ... Lendemer, Kocourkova & Knudsen
other lichenologists as the specimens bore no annotations by A.C.T.W. Herre
or J. Motyka who annotated many other Usnea specimens at PH. ‘The first
specimen in the folder was unidentified, however was immediately recognizable
as the pendant Eurasian arctic-boreal taxon Usnea diffracta which is widely
distributed in northwestern Asia (Japan, Korea, Siberia, Sakhalin Island, and
Taiwan (fide Motyka 1936-38)). Interestingly, the specimen had been collected
in Sitka, Alaska in 1898.
Sitka is a city on Baranof Island in the Alexander Archipelago in southeastern
Alaska, USA. The occurrence of Usnea diffracta in North America, especially
Alaska, is not unexpected considering the circumpolar nature of other Usnea
taxa such as U. filipendula Stirt. and U. longissima Ach. It is remarkable that
such a distinctive and conspicuous species has not been previously been
reported from North America (Esslinger 2008). The apparent lack of previous
records of U. diffracta in North America could simply be an indication of how
under-collected the continent is as a whole. While this explanation is a logical
one for the inconspicuous crustose taxa, cryptic taxa, and lichenicolous fungi
reported elsewhere in this paper it is unlikely that such a conspicuous lichen |
would have gone uncollected and unrecognized. Perhaps Usnea diffracta is a
rare species at the edge of its range in North America. Considering that the
collection reported here was made more than a century ago, in 1898, and no
further collections appear to have been mentioned in the literature, this is
likely the case. Hopefully this note will stimulate others to check their Usnea
collections for U. diffracta, and maybe additional collections, hopefully modern,
will be found. The lax pendant thallus, presence of annular pseudocyphellae
on the branches, and presence of diffractaic acid (CK+ yellow-gold, C-, K-)
in the medulla are diagnostic for U. diffracta. Descriptions and illustrations of
U. diffracta have been provided by Asahina (1956) and Ohmura (2001).
SPECIMEN EXAMINED. — U.S.A. ALASKA: Sitka, viii.1898, S.L. Schumo s.n. (PH).
Acknowledgements
Special thanks to Sarah Chaney of Channel Islands National Park, to Michaela Schmull of
the Farlow Cryptogamic Herbarium (FH), to Richard Harris of the New York Botanical
Garden (NY), Sean Beeching (Atlanta, USA), Leif Tibell (UPS), and Per Magnus
Jorgensen (BG). The work of J. Kocourkova was financially supported by a grant from
the Ministry of Culture of the Czech Republic (MK0000237201).
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MYCOTAXON
Volume 105, pp. 387-398 July-September 2008
Boletellus piakaii sp. nov. and a new distribution record for
Boletellus ananas var. ananas from Guyana
JORDAN R. Mayor’, TARA D. FULGENZzI’, TERRY W. HENKEL?
& Roy E. HALLING?
‘Department of Botany, University of Florida
Gainesville FL 32611 USA
*Department of Biological Sciences, Humboldt State University
Arcata CA 95521 USA
*Institute of Systematic Botany, The New York Botanical Garden
Bronx NY 10458 USA
Abstract — Boletellus piakaii (Boletaceae, Boletales, Basidiomycota) is described as
new to science. Boletellus ananas var. ananas is recorded for the first time from the
Guiana Shield region. These boletes were collected from tropical forests dominated
by ectomycorrhizal Dicymbe spp. (Caesalpiniaceae) in the Pakaraima Mountains of
western Guyana. A key is provided for Boletellus species known to occur in Guyana.
Key words — bolete, monodominant forest, tropical fungi, taxonomy
Introduction
The genus Boletellus Murrill (Boletaceae, Boletales, Basidiomycota) encompasses
~ 46 described species worldwide, the majority with tropical distributions
(Heinemann & Goossens-Fontana 1954, Snell & Dick 1970, Smith & Thiers
1971, Corner 1972, Horak 1977, Singer 1986 Singer et al. 1992, Watling 2001,
Halling & Mueller 2005, Ortiz-Santana et al. 2007, Fulgenzi et al. 2008). For a
discussion on various authors’ concepts of Boletellus see Fulgenzi et al. (2008).
Here we describe Boletellus piakaii sp. nov. and provide a new distribution record,
host association, and redescription for Boletellus ananas var. ananas occurring
in ectomycorrhizal (EM) Dicymbe (Caesalpiniaceae) forests of Guyana. The
new species is easily accommodated in Boletellus based on its olivaceous
brown, longitudinally ridged basidiospores, dry pileus, tubulose, blue-staining
hymenophore, and lack of clamp connections. Previous work in Guyana within
‘Corresponding author E-mail: twh5@humboldt.edu
388 ... Mayor & al.
this genus has lead to the description of two new species, Boletellus exiguus
T.W. Henkel & Fulgenzi and Boletellus dicymbophilus Fulgenzi & T.W. Henkel
(Fulgenzi et al. 2008). A key is provided for Boletellus taxa known to occur in
Guyana.
Materials and methods
Collecting expeditions were conducted during the May-July rainy seasons of
1997-99 in the Upper Ireng River Basin along Guyana’ western border with
Brazil in the west-central Pakaraima Mountains, and of 2000-07 in the Upper
Potaro River Basin ~ 30 km north of the Ireng River site. At each site fungi
were collected within a five km radius of a previously established base camp in
forests dominated by Dicymbe corymbosa Spruce ex Benth. (Henkel et al. 2002,
Henkel 2003).
Basidiomata were examined in the field for their fresh characteristics. Color
was subjectively described and recorded according to Kornerup & Wanscher
(1978) with color plates noted in parentheses (e.g., 3C4). Macro-chemical spot
tests were performed following Singer (1986). Basidiomata were field-dried ~
with silica gel beads.
Microscopic anatomical details were determined on fresh specimens at the
base camp with an EPOI microscope and in the laboratory using an Olympus
BX51 microscope with bright field and phase contrast optics. Fungal tissue
of dried specimens was rehydrated and mounted in either H 0; 3% KOH, or
Melzer’s solution. For each taxon a minimum of 20 basidiospores, basidia,
cystidia and other structures were measured. Line drawings were made with a
phototube, inked and scanned. Scanning electron micrographs of spores were
obtained with either a Jeol JSM-6400 or Topcon ABT32 scanning electron
microscope. Specimens are deposited in the following herbaria as noted: BRG,
HSU, and NY (Holmgren et al. 1990).
Taxonomic descriptions
Boletellus piakaii T.W. Henkel & Fulgenzi, sp. nov. FIGs 1-2
MycoBANK MB 511936
Pileus olivaceo-brunneus, rugosus et minute granulatus, 40-72 mm late; trama
caerulescens ubi exposita. Stipes stramineus, apice cum collo porphyreo, longitudinaliter
porcatus, 98-120 x 7-9 mm. Basidiosporae olivaceo-brunneae in massa, amygdaliformes,
longitudinaliter porcatae, 9.8-12.4 x 5-7.4 um.
Ho.LotyPe: Henkel 8728 (BRG; ISOTYPE: HSU, NY)
Erymo.oey. The species epithet recognizes the collector of the holotype, Piakai
Henkel.
KEY CHARACTERS — Boletellus piakaiiis recognized in the field by its olive-brown
rugose pileus, yellow-tan subequal stipe with maroon longitudinal striations,
Boletellus piakaii sp. nov. (Guyana) ... 389
Fig. 1. Basidioma of Boletellus piakaii (BRG HOLOTYPE Henkel 8728).
a. Dorsal. b. Ventral. Bar = 10 mm.
Fig. 2. Microscopic features of Boletellus piakaii (BRG HOLOTYPE Henkel 8728)
a. Basidia. b. Pleurocystidia. c. Basidiospores. Bar = 10 um.
390 ... Mayor & al.
and a rapid bluing reaction of the bruised hymenophore and exposed trama.
Microscopically B. piakaii is distinguished by having basidiospores that do not
exceed 12.5 um in length, basidia with long sterigmata and numerous golden
refractive globules, a cystoderm pileipellis of irregularly globose to subangular
cells, and numerous thick-walled hyphae in the pileus trama.
MACROCHARACTERS — PILEUS (25) 40-72 mm broad, convex to planoconvex,
dark brown (6F8) throughout when young to olive brown (5E6—-5E8—5F8)
to olive-yellow (4A8—4B8-—4C8-4D8) with age, rugulose to rugose, minutely
granular (under 10x lens), moist, margin slightly inrolled and entire; TRAMA
1-2 mm thick at margin, 3-5 mm over tubes, 6-12 mm above stipe, off-
white, immediately green-blue (26D5) upon exposure, fading to dingy-green
(28D6) and yellowing after several minutes, solid; TUBEs 3-6 mm long at
margin, 8-13 mm centrally, 6-7 mm at stipe, of irregular lengths, narrowly
and deeply depressed around stipe, with short decurrent teeth 1-2 mm long,
light olive-yellow (3A7—3B7), quickly blue-green (20F8—24F8) upon exposure,
eventually fading to dull brown; porzs yellow (2A8) when immature to olive-
yellow (3A7-3B7-4B8-4C8), immediately blue-green (24F8) with pressure,
then fading gradually to green (27F8), green-brown (4E6), and eventually
brown (6E8-6F8), 1-1.5 per mm, subisodiametric to angular; sTIPE (60)
98-120 x 7-13 mm, subequal, enlarging gradually to 11-13 mm broad at
base, ground tissue yellow-tan (4A5-4B5-—4C8), overlaid with pinkish brown
(6D5-6E5-6D4-6F4) to red-maroon (7E6-7E7-7D8-8E7) longitudinal
striations throughout, these darkening with age and handling, apex with light
red-brown (7A5) band, lower 20-25 mm with densely matted dark-brown
(7F4—-8F2) tomentum; SUBTENDING HYPHAE con-colorous and often associated
with ectomycorrhizal rootlets; TRAMA off-white to yellow (4A7-4B7) to brown
(7D5-—7E5-7D8-8E7) with age, transiently bluing upon exposure, solid; ODOR
minimal, pleasant; TASTE mild.
MICROCHARACTERS — BASIDIOSPORES dark olive-brown (4F8—5F8) in
medium to heavy deposit, 9.8-12.4 x 5-7.4 um (mean Q = 1.69), golden-
olive in H,O and KOH, inamyloid, amygdaliform, 1-2 guttulate, with 18-22
longitudinal ridges; ridges < 1 um tall, repeatedly dichotomously forked,
converging at both poles, transverse striae absent; hilar appendage 0.5-0.9
um long; BASIDIA 32.4—46 x 13.1-15.6 um, clavate, wall hyaline in H,O and
KOH, with numerous refractive globules, these light golden in H,O, lighter
in KOH, 4-sterigmate; STERIGMATA 3.2—5 um long; PLEUROCYSTIDIA 37-69
x 8.4-12.4 tm, ventricose-rostrate, frequent, arising from the subhymenium,
projecting 21-30.8 um beyond the hymenial palisade, hyaline, thin-walled;
CHEILOCYSTIDIA absent; HYMENOPHORAL TRAMA boletoid, strongly divergent;
MEDIOSTRATUM 19.7—49.4 tum wide, of many thin-walled, nearly parallel
hyphae, golden in H,O, lighter and gelatinized in KOH; LATERAL STRATUM
Boletellus piakaii sp. nov. (Guyana) ... 391
hyphae 7.4—12.4 ttm wide, hyaline, nongelatinized; PILEIPELLIS a cystoderm of
irregularly globose to subangular cells; these thin-walled, faint-golden in H,O,
hyaline in KOH; piLeus TRAMA of interwoven hyphae, many with thickened
walls; individual hyphae 4.4-11.1 «um wide, wall up to 0.5 um thick, regularly
septate, devoid of contents, in mass light golden in H,O, lighter in KOH;
STIPITIPELLIS an interwoven trichodermial palisade of cylindrical elements
with inflated terminal cells, in mass golden orange in H,O, lighter in KOH;
TERMINAL CELLS projecting 22.5-40 um, cylindrical to clavate, occasionally
ventricose to mucronate, these concentrated on longitudinal striations;
SUBPELLIS hyphae interspersed with external burgundy acerose crystals; STIPE
TRAMA Of densely packed parallel hyphae, in mass faint golden in H,O, lighter
in KOH; individual hyphae 4—9.6 um wide, hyaline in H,O and KOH, regularly
septate; CLAMP CONNECTIONS absent.
MACROCHEMICAL REACTIONS — NH,OH rapidly light red-brown (8D6) on
pileus, soon darkening, burgundy (9E8) on stipe ground, black on longitudinal
striations, light burgundy (9D6) on stipe trama; KOH light orange-brown
(7C6) to burgundy (8F7) on pileus, burgundy (8F7) on stipe, red-brown (8D8)
on pileus and stipe trama; FeSO, dark grey-green (28F4) on pileus, dark green
(29F5) on tube mouths, dark green (28F8) on stipe, yellow-green (29D6) on
stipe trama.
ECOLOGY, RANGE, DISTRIBUTION — Solitary to scattered on root mat in
D. corymbosa forest, rarely on lower base of D. corymbosa trunks; more frequent
late in the rainy season; known only from the type locality in the Upper Potaro
River Basin of Guyana.
REPRESENTATIVE SPECIMENS EXAMINED. GUYANA. REGION 8: POTARO-SIPARUNI.
Pakaraima Mountains, Upper Potaro River, within a 5 km radius of ‘Potaro Base Camp’
(5° 18’ 04.8” N, 59° 54’ 40.4” W, elevation 710-750 m); vicinity of Potaro base camp, 2
May 2001, Henkel 8013 (BRG; HSU); in Dicymbe plot 1, 9 May 2001, Henkel 8077 (BRG;
HSU), 2 July 2004, Henkel 8728 (HOLOTYPE, BRG; ISOTYPE: HSU, NY); 1 km west of
Potaro base camp, 30 May 2005, Henkel 8810 (BRG; HSU); 1.5 km southeast of Potaro
base camp on ‘Benny’s ridge, 14 August 2007, Henkel 8869 (BRG; HSU); in Dicymbe plot
2, 24 August 2007, Henkel 8878 (BRG; HSU); 0.75 km northeast of Potaro base camp
between north end of “Benny’s ridge’ and ‘Leon camp, 25 August 2007, Henkel 8882
(BRG; HSU).
COMMENTS — Within the genus B. piakaii is best disposed in Section
Chrysenteroidei Singer due to the longitudinally ridged basidiospores, dry
pileus lacking in red tints, and the absence of a veil (Snell & Dick 1970, Singer
1986). Boletellus piakaii is similar in stature and shares the brown, rugose pileus
reported for Boletellus longicollis (Ces.) Pegler & T.W.K. Young from Malaysiaand
Japan; B. longicollis can be distinguished by its white, gelatinous, appendiculate
veil and non-cyanescent trama (Corner 1972, Imazeki et al. 1988). Among
other Malaysian boletelli only Boletellus fallax (Corner) Watling non Singer has
392 ... Mayor & al.
cyanescent trama and lacks an appendiculate veil; B. fallax otherwise differs
from B. piakaii in having longer basidiospores (18-24 um vs. 9.8-12.4 um),
longer pleurocystidia (60-140 um vs. 37-69 um), a smaller stature (pileus diam
25-45 mm vs. (25) 40-72 mm, stipe length 22-70 mm vs. (60) 98-120 mm),
and red tints in the pileus (Corner 1972). Among Congolian Boletellus species
none are similar to B. piakaii; only Boletellus longipes Heinem. has a similar
stature, but larger basidiospores (12.7-16.2 x 7.2-8.5 um vs. 9.8-12.4 x 5-7.4
lum), an orange stipe, and areolate pileus (Heinemann & Goossens-Fontana
1954). The Central American Boletellus belizensis B. Ortiz & T.J. Baroni and
Boletellus domingensis B. Ortiz & Lodge lack marginal appendiculae and have
a cyanescent reaction in the exposed pileus trama and bruised hymenophore,
yet these species are dissimilar to B. piakaii in having smaller statures (pileus
diam 24—40 and 18-60 mm vs. (25) 40-72 mm, stipe length 60-70 and 27-55
mm vs. (60) 98-120 mm), larger basidiospores (12-14.4 x 5.6-7.2 and 12.8-16
x 5.6-8 um vs. 9.8-12.4 x 5-7.4 um) pruinose stipe ornamentations, and in
lacking a rugulose to rugose pileus surface (Ortiz-Santana et al. 2007).
Boletellus ananas (M.A. Curtis) Murrill, Mycologia 1:10. 1909 var. ananas
FIGS 3-5
= Boletus ananas M.A. Curtis Amer. J. Sci. Arts 6: 351. 1848.
= Boletus coccineus Fr. Epic. Myc. 423. 1838, nom. illegit., non Bull. 1791.
= Strobilomyces coccineus Sacc., Syll. Fungorum 6: 50 1888 ut “Fr”.
= Boletellus coccineus (Sacc.) Singer in Singer et al. Beih. Nova Hedwigia 105: 6: 1992
ut “(Fr)”
= Boletus isabellinus Peck, Bull. Torrey Bot. Club 24: 146. 1897, nom. illegit., non Schwein.
1322)
= Strobilomyces pallescens Cooke & Massee in Cooke, Grevillea 18: 5 1889.
= Boletellus pallescens (Cooke & Massee) E.-J. Gilbert, Bolets. 107. 1931.
KEY CHARACTERS — Boletellus ananas var. ananas is recorded for the first time
from northeastern South America. This striking bolete is recognized in the field
by its red-pink squamose pileus, membranous marginal appendiculae, bright
yellow, rapidly blue-staining hymenophore, cyanescent trama, and tree trunk
fruiting habit. These features are consistent with those of B. ananas var. ananas
as described from other regions (McNabb 1967, Smith & Thiers 1971, Corner
1972, Singer 1986, Halling & Mueller 2005). Microscopically B. ananas var.
ananas is distinguished by large basidiospores with cross striae on the ridges
and spirally encrusted hyphae in the marginal appendiculae and stipe trama.
MACROCHARACTERS — PILEUS 33-74 mm wide, convex to planoconvex,
with appressed to recurved squamules, red-brown (10B7—10D7) to red-tan
(7D5-8D4-8E5) to pink (9A4—10B4) to pink-grey(10C4), vestiture concentrated
and more squarrose on disk, extending out of cream (1A2—3A2-3A3-4A3)
to light orange-pink (7A3) to light pink-red (9A6) floccose ground; margin
Boletellus piakaii sp. nov. (Guyana) ... 393
Fig. 3. Basidiomata of Boletellus ananas var. ananas.
a.Hsu Henkel 8819. b. Fruiting habit on humic deposit of Dicymbe trunk, Upper Potaro Basin,
Guyana. Bar = 10 mm.
Fig. 4. Microscopic features of Boletellus ananas var. ananas (Hsu Henkel 8168)
a. Basidia. b. Pleurocystidia. c. Basidiospores. Bar = 10 um.
entire and clasping stipe when young, at maturity separating into triangular
appendiculae, these 6-12 x 3-10 mm, buff white (5A3-5B4), occasionally faint
pink; moist; trama 2-3 mm thick at margin, 7-10 mm over tubes, 11-18 mm
centrally, buff white to light yellow (3A3), rapidly bluing (23C7) upon exposure,
solid; ruBEs 1-5 mm long at margin, 10-20 mm centrally, 4-6 mm at stipe,
broadly and deeply depressed around stipe, of irregular lengths, bright yellow
394 ... Mayor & al.
(3A8) to olive yellow (2B7-3B7-3C7) to mustard yellow (4A8-4B8), rapidly
bluing (23C7).upon exposure; PORES concolorous with tubes, rapidly blue-green
(23C8) with pressure, 0.5-1.5 per mm, angular; sTIPE 53-115 x 6-14 mm broad,
subequal, gradually enlarging toward base to 10-19 mm, apical portion cream
(3A4) to pink (9A4), middle portion finely longitudinally striate, striations
darkening with handling, red-lavender (11A6-11B4-11B7-11B8) to brown-
red (9E8-10E8), lighter with age, immediately above basal tomentum cream
(3A2-3A4) with few striations; BASAL TOMENTUM of white strigose hairs over
lower 6-50 mm, occasionally subtended by ectomycorrhizal rootlets; TRAMA
white to buff-tan (5B4) to light yellow (3A4), slightly bluing with exposure;
solid; opoR non-descript; TASTE mild.
MICROCHARACTERS — BASIDIOSPORES Olivaceous-brown (4D7-4E7-5E6—5F6)
in medium to heavy deposit, 17.5-22.2 x 6.4-8 um (mean Q = 2.73), golden
in H,O and KOH, inamyloid, amygdaliform, one to multiguttulate; wall 0.5—-1
um thick, with 12-14 longitudinal ridges; ridges < 1 um tall, occasionally
bifurcating, converging at poles, with minute cross-striae; hilar appendage
0.3-1 um long; Basrp1A 39-57 x 11-15 um, clavate, wall hyaline in H,O and. |
KOH, with numerous refractive globules, these faint golden in H,O and KOH,
4-sterigmate; PLEUROCYSTIDIA 42-47 x 8-12 um, ventricose-rostrate, slightly
capitate, frequent, arising from the sub-hymenium, projecting 19.3-29.6
uum above the hymenial palisade, thin-walled, hyaline, devoid of refractive
contents; CHEILOCYSTIDIA 19-42 (51) x 5-11 um, ventricose, cylindrical, to
narrowly clavate, thin-walled, infrequent; HYMENOPHORAL TRAMA boletoid,
strongly divergent; medio-stratum 24.7-45.7 um wide, of many parallel,
slightly interwoven hyphae, faintly grey-yellow in H,O, lighter and gelatinized
in KOH; lateral stratum hyphae 4.4-8.4 um wide, hyaline and gelatinized
in KOH, regularly septate; PILEIPELLIS a densely interwoven trichodermial
palisade of cylindrical elements with inflated terminal cells, in mass faint
golden in H,O, lighter in KOH; terminal cells 23.5-51.9 x 9.4-16.8 um, hyaline
in H,O and KOH, inamyloid, cylindrical to clavate, interwoven, concentrated
on squamules; marginal appendiculae composed of wefts of interwoven inflated
hyphae, some with faint golden spirally arranged encrusting pigments evident in
HO, KOH, and Melzer’s; pILEUS TRAMA highly interwoven; individual hyphae
7.4-11.1 um wide, hyaline in H,O, gelatinized and hyaline in KOH, regularly
septate; STIPITIPELLIS a trichodermial palisade of cylindrical elements with
inflated terminal cells, in mass golden in H,O, lighter in KOH; terminal cells
projecting 30.4—63 um, cylindrical to clavate, occasionally mucronate; STIPE
TRAMA densely interwoven, in mass golden in H,O, lighter in KOH; individual
hyphae 4.9—7.2 um wide, hyaline in H,O and KOH, with spirally arranged faint
golden encrusting pigments, these evident in KOH, Meltzer’s, and H,O; CLAMP
CONNECTIONS absent.
Boletellus piakaii sp. nov. (Guyana) ... 395
Fig. 5. Scanning electron micrographs of basidiospores of Boletellus species from Guyana.
a. Boletellus piakaii (Hsu Henkel 8013) x4000.
b. Boletellus ananas var. ananas (HsU Henkel 8168) x6300.
MACROCHEMICAL REACTIONS — NH,OH olivaceous yellow (4D6) on pileus,
dirty yellow (4C7) on tube mouths, bright red (10B8) on stipe, tan (6C4) on
basal tomentum, yellow on trama of pileus and stipe; KOH bleaching pileus
ground, rusty-brown (7D8) on squamules, orange-brown (5C6) on stipe,
orange-yellow on pileus trama, burgundy to orange on stipe trama; FeSO, pink
to faint grey on pileus, steel-grey on stipe; olive-green (26F6) on tube edges.
ECOLOGY, RANGE, DISTRIBUTION — In Guyana typically fruiting singly or in
pairs within 1-2 m above ground level on D. corymbosa trunks, associated with
ectomycorrhizas within humic accumulations; rarely found fruiting on ground
on heavily decayed, root-penetrated wood; recorded from the Upper Potaro
and Upper Ireng River Basins in Guyana; also known from south-eastern North
America, Central America, New Zealand, Malaysia, and possibly Australia
(Corner 1972, Singer et al. 1983).
REPRESENTATIVE SPECIMENS EXAMINED — BRAZIL, AMAZONAS: Boletellus ananas var.
minor, Estrada Manaus-Caracarai, 4 April 1980, Singer B10913 (HOLOTYPE, INPA).
GUYANA, REGION 8, Potaro-Siparuni, Pakaraima Mountains, Upper Ireng River Basin,
west bank of Suruwabaru Creek, 1 km upstream from confluence with Yuarka creek, 2
March 1997, Henkel 6263- 6264; 23 May 1999, Henkel 7038; Upper Potaro River, within
a5 km radius of ‘Potaro Base Camp’ (5° 18’ 04.8” N, 59° 54’ 40.4” W, elevation 710-750
m); Ayanganna airstrip, 19 May 2000, Henkel 7400; vicinity of Potaro base camp, 30
June 2000, Henkel 7925; 4 km southwest of Potaro base camp near Dicymbe plot 3, 18
May 2001, Henkel 8168; vicinity of Potaro base camp, 25 July 2003, Henkel 8614, 5 July
2004, Henkel 8734, 31 May 2005, Henkel 8819; 2 km south of Potaro base camp, 2 June
2005, Henkel 8826; 0.75 km west of Potaro base camp, 3 June 2005, Henkel 8833. Guyana
collections housed in BRG and HSU.
396 ... Mayor & al.
COMMENTARY — We have not adopted the nomenclature erected by Singer
et al. (1992: ice. Boletellus coccineus var. coccineus for Boletellus ananas var.
ananas). Singer created this name in the mistaken belief that the earliest
available name for the taxon was Boletus coccineus Fr. 1838. However, Fries’s
name is an illegitimate later homonym (cf. Boletus coccineus Bull. 1791), and
Singer's combination is actually based on Strobilomyces coccineus Sacc. 1888.
The earliest available name for the taxon is therefore Boletus ananas M.A.
Curtis 1848, the basionym of Boletellus ananas.
Boletellus ananas var. ananas, as the type of the genus, is in Section Boletellus
Singer based on the squamose, dry pileus with red-pink tones, a marginal veil
that clasps the stipe when immature, and longitudinally ridged spores that are
> 16 um long (Singer 1986). Most characters of the Guyana material other than
host association are consistent with those previously reported for B. ananas var.
ananas; one minor deviation is that the pleuro-cystidia of Guyana specimens
are consistently shorter (< 50 um) than those reported for collections from
Malaysia (50-95 um), and the Dominican Republic and Belize (44-91.2 um)
(Corner 1972, Ortiz-Santana et al. 2007), but consistent with those of material
from southeastern North America, New Zealand, and Costa Rica (McNabb
1967, Smith & Thiers 1971, Singer et al. 1983). Boletellus ananas var. ananas
has consistently larger basidiomata than B. ananas var. minor Singer from
Brazil and Nicaragua, and lacks the thick-walled cheilocystidia of B. ananas
var. crassotunicatus Singer from Nicaragua and Panama and laciniate-reticulate
stipe ornamentation of B. coccineus var. amarus Singer from Honduras and
Belize (Singer et al. 1983, Singer et al. 1992, Ortiz-Santana et al. 2007).
The habit of fruiting on tree trunks has been reported for varieties of
B. ananas from southeastern North America, Costa Rica, Brazil, Panama,
Nicaragua, and Guyana (Murrill 1909, Singer et al. 1983, this paper), although
terrestrial fruiting has been reported in Malaysia (Corner 1972) and Central
America (T. Baroni pers. comm.). Due to the typically elevated fruiting habit
and occurrence on dead wood the ectomycorrhizal status of B. ananas has been
debated, although all collections have been made in association with ectotrophic
host trees including Pinus and Quercus spp. in southeastern North America
(Murrill 1909) and Central America (Singer et al. 1983, Halling & Mueller 2005,
Ortiz-Santana et al. 2007), Quercus humboldtii Bonpl. in Colombia (Franco-
Molano & Uribe-Calle 2000), various Fagaceae and Dipterocarpaceae spp. in
Malaysia (Corner 1972), and Leptospermum and Pinus spp. in New Zealand —
(McNabb 1967). In Guyana the humic deposits on Dicymbe trunks bearing
B. ananas are consistently permeated with abundant ectomycorrhizas (Woolley
et al. 2008).
=
a
Boletellus piakaii sp. nov. (Guyana) ... 397
Boletellus in Guyana
Key to Boletellus species
la. Marginal veil present, becoming appendiculate; pileus squamose to
squamulose, with pinkish tonesee rn eh eaniiad Jechias: 20) B. ananas var. ananas
1b. Marginal veil absent; pileus squamulose or otherwise, lacking pinkish tones .... 2
2a. Fruiting terrestrial, rarely on tree trunks; pileus > 46 mm broad;
stipe > 98 mm long; basidiospores not exceeding 12.5 7um........ B. piakaii
2b. Fruiting on tree trunks; pileus < 45 mm broad; stipe < 80 mm long;
asi COSDOLES UP {Osho < aL lr lieben ter Oe ee ee epee a ee a 3
3a. Basidiomata small; pileus 9-33 mm broad; stipe 7-34 x 1-4 mm;
pleurocystidia infrequent, ventricose, 65-85 (100) x 14-20 um ..... B. exiguus
3b. Basidiomata larger; pileus 19-34 mm broad; stipe 41-60 (80) x
4—7 mm; pleurocystidia frequent, cylindrical to narrowly clavate,
BB SOS IDS AO RR ree est B. dicymbophilus
Acknowledgements
This research was supported by grants to TWH from the National Geographic Society's
Committee for Research and Exploration, the Smithsonian Institution’s Biological
Diversity of the Guiana Shield Program, the Linnaean Society of London, the Duke
University Department of Botany, and the Humboldt State University Foundation. Field
assistance in Guyana was provided by Mimi Chin, Cathie Aime, Christopher Andrew,
Leonard Williams, Valentino Joseph, Francino Edmond, and Luciano Edmond. REH
acknowledges support from the National Science Foundation grant DEB-0414665. The
authors would like to thank J.W. Kimbrough and G. Laursen for use of laboratory facilities,
and Christian Feuillet for the Latin translations. Nomenclatural clarifications from Scott
Redhead were greatly appreciated. Excellent critical reviews of the manuscript were
provided by boletologists Beatriz Ortiz-Santana and Timothy Baroni. Shaun Pennycook
provided valuable nomenclatural advice that greatly improved the manuscript. Research
permits were granted by the Guyana Environmental Protection Agency and Ministry
for Amerindian Affairs. This paper is number 131 in the Smithsonian Institution's
Biological Diversity of the Guiana Shield Program publication series.
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MYCOTAXON
Volume 105, pp. 399-405 July-September 2008
New records of Scleroderma from Northeastern Brazil
EE. GurGEL, B.D.B. SILVA & I.G. BASEIA
baseia@pesquisador.cnpq. br
Universidade Federal do Rio Grande do Norte, CB,
Depto. de Botanica, Ecologia e Zoologia
CEP: 59072-970, Natal, RN, Brazil
Abstract — Survey on species of Scleroderma in native trees from the Atlantic rain
forest biome, and introduced cultures of Pinus, was performed in the States of Paraiba,
Pernambuco and Rio Grande do Norte. Scleroderma albidum, S. bovista, S. citrinum and
S. nitidum were identified, the first three being new records to the Northeast Brazil and
the latter, to the country. Descriptions of macro and microscopic features and SEM-
photos of the spore ornamentation of these species are given.
Key words — gasteromycetes, Boletales, taxonomy, biodiversity
Introduction
The genus Scleroderma includes 25 ectomycorrhizal taxa usually known as earth
balls, distributed worldwide (Kirk et al. 2001). This genus of poisonous fungi
with hard-skinned basidiomata is recognized by its epigeous and single-layered
peridium opening by irregular dehiscence and gleba without capillitium.
To date, Scleroderma albidum, S. areolatum Ehrenb., S. bougheri Trappe, et al.,
S. bovista, S. cepa Pers., S. citrinum, S. floridanum Guzman, S. fuscum (Corda)
E. Fisch., S. uruguayense (Guzman) Guzman, S. verrucosum (Bull.) Pers.,
S. vulgare Hornem., S. tenerum Berk. & M.A. Curtis and S. polyrhizum (J.E Gmel.)
Pers. have been reported in Brazil by Rick (1961), Guzman (1970), Bononi et
al. (1981), Baseia & Milanez (2000) and Giachini et al. (2000). These species
have been investigated for their anti-inflammatory and hemostatic properties
(Guzman 1970, Liu 1984). In nature, they are important ectomycorrhizal fungi
and many species have been studied in light of their potential to enhance tree
growth (Kropp & Trappe 1982, Molina & Trappe 1982, Danielson 1984, Garrido
1984, Jones et al. 1986, Richter & Bruhn 1989, 1990; Richter 1992). This paper
reports the species of Scleroderma found in native forests from the Atlantic
rainforest biome and in introduced cultures of Pinus, the main purpose being
to increase the knowledge about this taxon.
A(0 ... Gurgel, Silva & Baseia
Material and methods
Material was colleted during field trips to Atlantic rainforest of the States of
Paraiba, Pernambuco and Rio Grande do Norte. The material was deposited
in the Herbarium of the Universidad of Pernambuco, Brazil (URM), and in
the Herbarium of the Universidade Federal do Rio Grande do Norte, Brazil
(UFRN-fungos).
Hand-cut sections of dried material were mounted in Melzer’s reagent,
5% KOH, water or cotton blue for microscopic examination (Guzman 1970).
For SEM pictures, the spores were air-dried, mounted on aluminum stubs,
sputter coated (Bal-Tec SCD-050) with gold and examined in a Philips XL 20
scanning electron microscope. The mycorrhizal observations were performed
using methods developed by Agerer (1991) and Fortin (1980). Color terms in
parenthesis are those of Kornerup & Wanscher (1978).
Results
Scleroderma albidum Pat. & Trab., Bull. Soc. Myc. Fr. 15: 57, 1899. FIGS 1, 5.
Basidiomata 2-4 cm wide, subglobose or depressed globose, yellowish white
(4A4) with a small rhizomorphic base. Peridium up to 5 mm thick when
fresh, becoming thinner when dry, smooth surface, usually dehisced by
irregular splitting, from the apex downwards into unequal lobes; branched or
unbranched thin walled hyphae, 3-4 um diam. Gleba pulverulent, dark grey
(3F3). Basidiospore globose, echinulate, 12-14 um diam, dark brown.
HasitTatT: On sandy soil, epigeous, aggregated with roots of Pinus radiata
D. Don in a plantation of this tree.
MATERIAL EXAMINED: BRAZIL. PERNAMBUCO: RECIFE, HOoRTO FLORESTAL Dols
IrMAos, 18.VII.2007, I.G. Baseia (URM 307577).
ADDITIONAL MATERIAL EXAMINED: BRAZIL, Sao Paulo, Campos do Jordao, Parque
Florestal, 09/III/1971, G. Guzman, SP 214722; SP 112060.
DISTRIBUTION: Brazil (Guzman 1970, Bononi et al. 1981, Giachini et al. 2000), Mexico
(Esqueda- Valle et al. 1990).
COMMENTS— Scleroderma albidum is characterized by its light brown and
smooth peridial wall. It differs from S. polyrhizum, which is recognised by the
rough surface of the peridium and subglobose spores with smaller spines (Baseia
& Milanez 2000). The basidiomes of the material studied were found growing
alone or in groups during the rainy season. S. albidum was found associated
with Pinus radiata, an exotic conifer from North America introduced into some
Atlantic rainforest areas. Its basidiomes presented rhizomorphs in aggregation
with the roots of this tree. The root tips showed typical ectomycorrhiza with
mantle and Hartig net. It is reported for the first time in Northeastern Brazil.
|
»
EL
Scleroderma spp. new to Brazil ... 401
=
io)
ie
Co
&
oO
PP
aD,
Fig. 1-4. Basidiomata:
1. Scleroderma albidum; 2. S. bovista; 3. S. citrinum; 4. S. nitidum
Scleroderma bovista Fr., Syst. Myc. 3: 48, 1829. FIGS 2, 6
Basidiomata 2-3 cm wide, subglobose, irregularly globose to oblate, yellowish
brown (5E5). Peridium up to 4-6 mm thick when fresh, becoming thinner
when dry, smooth surface, usually dehisced by irregular splitting; thin walled
hyphae, unbranched, 3-4 um diam, clamp connections present. Gleba compact,
yellowish brown (6F7). Basidiospore brown, globose, echinulate and reticulate,
9-11 um diam.
Hasitat: On sandy soil, epigeous, aggregated with roots of Gomidesia spectabilis
(DC.) O. Berg (Myrtaceae), in Atlantic rainforest.
MATERIAL EXAMINED: BRAZIL. PERNAMBUCO: HORTO FLORESTAL Dols IRMAOs,
29.VII.2007, I.G. Baseia (UFRN-fungos 456).
ADDITIONAL MATERIAL EXAMINED: BRAZIL, Sao Paulo, Campos do Jordao, Parque
Florestal, 09/III/1971, G. Guzman, SP 214720.
DISTRIBUTION: Argentina (Spegazzini 1927), USA (Coker & Couch 1928, Richter 1992),
South Africa (Bottomley 1948), Norway (Eckblad 1955), Congo (Dissing & Lange 1962),
China (Liu 1984), Chile (Garrido 1986), Brazil (Guzman 1970, Giachini et al. 2000).
402 ... Gurgel, Silva & Baseia
CoMMENTS— Scleroderma bovista is recognised by its smooth peridial surface,
differing from S. albidum mainly by the basidiospores, which are smaller and
reticulate in S. bovista. The basidiomata of the material studied were found
growing alone or in groups during the rainy seasons associated with Gomidesia
spectabilis, an ectomycorrhizal tree native to this region. The symbiotic
relationship is sustained by rhizomorphs of S. bovista in aggregation with the
roots of this tree. A mantle and Hartig net, characteristics of ectomycorrhizae,
were detected by microscopic analysis. The species is thought to have a
wide host range (Guzman 1970): Garrido (1986) noted an ectomycorrhizal
association between this fungus and Pinus radiata in Chile and Giachini et al.
(2000) noted association with Pinus and Eucalyptus species. This is the first
report of S. bovista in Northeastern Brazil.
Scleroderma citrinum Pers., Syn. Meth. Fung. 1: 153, 1801. FIGS 3, 7
Basidiomata, light yellow (4A5), depressed globose, 4-7 cm wide, attached to
the soil by a small rhizoidal base. Peridium up to 2-3 mm thick when fresh,
surface areolate, dehisced by irregular cracking of the apex; hyphae thin walled, |
usually unbranched, 2-3 um diam, clamp connections present. Gleba compact,
becoming pulverulent, yellowish brown (5F6). Basidiospores yellowish brown,
globose or subglobose, echinulate and reticulate, 12-13 um diam.
HasitTat: On sandy soil, epigeous in a Pinus taeda L. plantation, aggregated
with roots of this tree.
MATERIAL EXAMINED: BRAZIL. PARAIBA: JOAO PESSOA, JARDIM BOTANICO,
11.V1.2006, I.G. Baseia (UFRN-fungos 502).
ADDITIONAL MATERIAL EXAMINED: USA, Maryland, 21/VH/1958, G. Guzman, SP
50636.
DISTRIBUTION: Spain (Calonge & Demoulin 1975), Brazil (Guzman 1970, Bononi et
al. 1981, Giachini et al. 2000), China (Liu 1984), Chile (Garrido 1986), USA (Richter
1992).
COMMENTS— Scleroderma citrinum is characterized by its large yellowish white
basidiomata and peridium with areolate surface. The basidiomata were found
growing abundantly during the rainy seasons in ectomycorrhizal association
with Pinus taeda. Garrido (1986) found an ectomycorrhizal association between
this fungus and Pinus radiata in Chile, demonstrated by pure culture synthesis.
During our fieldwork we observed that S. citrinum was the most common
Scleroderma species in the pine plantation. This is the first report of this fungus
in Northeastern Brazil.
Scleroderma nitidum Berk., Hook J. Bot. 6: 173, 1854. Fics 4, 8
Basidiomata yellowish brown (5D5), subglobose, 1.5-3 cm wide. Peridium up to
4—5 mm thick when fresh, becoming thinner when dry, verrucose surface, dark
Scleroderma spp. new to Brazil ... 403
Figs. 5-8. SEM pictures of basidiospores:
5. Scleroderma albidum; 6. S. bovista; 7. S. citrinum; 8. S. nitidum
brown (6F6), evenly scattered on a surface that dehisced by irregular splitting;
hyphae thin-walled, hyaline, 10-18 x 6-12 um. Gleba pulverulent, olive grey
(3F3). Basidiospore dark brown, globose, echinulate, 11-12 um diam.
Hasirat: On sandy soil, epigeous under Coccoloba sp.
MATERIAL EXAMINED: BRAZIL. PARA{BA: JOAO PESSOA, JARDIM BOTANICO,
24.VI1I.2007, I.G. Baseia (UFRN-fungos 464). PERNAMBUCO: RECIFE, HORTO FLORESTAL
Dois IRMAos, 15.V1.2007, LG. Baseia (UFRN-fungos 494). BAHIA: SENHOR DO BOFIM,
17.111.2006, I. Gusmao (UFRN-fungos 500). R10 GRANDE DO NorTE: NATAL, PARQUE
DAs Dunas, 03.V.2007, EE. Gurgel (UFRN-fungos 491).
ADDITIONAL MATERIAL EXAMINED: USA, Maryland, VII/1958, D. M. Dring, SP 55938;
AFRICA, SIERRA LEONE, 22/V1/1965, G. Guzman, SP 107416.
DIsTRIBUTION: Argentina (Spegazzini 1927), South Africa (Bottomley 1948), Norway
(Eckblad 1955), Panama (Garner 1956), Brazil (Rick 1961, Guzman 1970, Giachini et al.
2000), Congo (Dissing & Lange 1962), Spain (Calonge & Demoulin 1975), China (Liu
1984), Chile (Garrido 1986), Mexico (Pardavé 1991), Belgium (Moyersoen & Demoulin
1996).
COMMENTS — This species is usually easy to distinguish by its yellowish brown
and soft peridium, covered by dark brown verrucae at least over the upper
part,. It is close to S. citrinum but differing in the colour and consistency of
the peridium. The basidiomata were found growing in groups during the rainy
404 ... Gurgel, Silva & Baseia
seasons in ectomycorrhizal association with the native tree, Coccoloba sp. Its
roots tips showed typical ectomycorrhiza with mantle and Hartig net. This is
the first report of S. nitidum in Brazil. 3
Acknowledgments
We express our gratitude to the Conselho Nacional de Desenvolvimento Cientifico e
Tecnoldgico (CNPq), for the financial support and to Tereza Cristina de Oliveira Galvao
for the illustrations. We also thank Professors Leticia Montoya and Tatiana Baptista
Gilbertoni for critical revision of the manuscript.
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MYCOTAXON
Volume 105, pp. 407-414 July-September 2008
Description of Tuber malenconii mycorrhizae
and a comparison with mycorrhizae of
T. aestivum and T. mesentericum
Luis G. GARCIA-MONTERO'™, GABRIELLA D1 MASSIMO’,
JosE L. MANJON? & JAVIER VELAZQUEZ4
luisgonzaga.garcia@upm.es
‘Dept. Forest Engineering, U.D. Operaciones Basicas, E.T.S.I. Montes
Technical University of Madrid (UPM)
Ciudad Universitaria s/n, Madrid 28040, Spain
"Dept. di Biologia Vegetale, Universita degli studi di Perugia
Borgo XX Giugno 74, 06121 Perugia, Italy
*Dept. Biologia Vegetal, Sciences Faculty, Universidad de Alcala
28871 Alcala de Henares (Madrid), Spain
“Dept. of Projects and Rural Planning, E.T:S.I. Montes,
Technical University of Madrid (UPM)
Ciudad Universitaria s/n, Madrid 28040, Spain
Abstract — Tuber malenconii is a recently described uncommon European truffle
associated with Quercus ilex that is potentially detrimental to the cultivation of the
highly prized edible, T’ melanosporum. Examination of T: malenconii mycorrhizae
synthesized from fresh ascocarps revealed polygonal mantle cells and numerous cystidia
with basal-forked ramifications. Differences between T: malenconii, T. mesentericum,
and T: aestivum mycorrhizae are noted; compound and scanning electron micrographs
provide additional comparisons.
Key words — Tuberaceae, ectomycorrhizae, hypogeous fungus
Introduction
The recently described truffle, Tuber malenconii, is a rather rare species that has
been reported from France (Donadini et al. 1979, Riousset et al. 2001), Italy
(Montecchi & Sarasini 2000), and Spain (Alvarez et al. 1993). T; malenconii
inhabits warm regions in clay brown calcareous soils under Quercus ilex L.
This truffle has an unpleasant faecal odour and therefore no economic value.
The fact that T: malenconii has been collected in cultivations of the highly
prized black truffle, T; melanosporum Vittad., in France and Spain suggests
A408 ... Garcia-Montero & al.
that it is a potential competitor that might be detrimental to T’ melanosporum
carpophore production. Garcia-Montero et al. (1996) and Manjon & Garcia-
Montero (1996) have cited the synthesis of T: malenconii mycorrhizae and have
expressed concern for the risk that this species would pose to T: melanosporum
should it be introduced (accidentally or otherwise) into truffle plantations.
It is the usual practice to carry out quality control in nurseries that
produce mycorrhized plants for truffle culture. Several authors have detailed
the morphology of the mycorrhizae of economically valuable Tuber species
(Zambonelli et al. 1993, Granetti 1995). However, in order to establish efficient
quality controls of mycorrhized plants, we need to know more about all
Tuber mycorrhizae, as demonstrated in the comparisons between the similar
ectomycorrhizae representing some Chinese Tuber species and T’ melanosporum
(Di Massimo et al. 1996, Comandini & Pacioni 1997, Zambonelli et al. 1997,
Manjon et al. 1998, Garcia-Montero et al. 2008). Accordingly, we describe the
morphology of T: malenconii ectomycorrhizae, which we then compare with
the similar mycorrhizae of T: aestivum Vittad. and T. mesenterica Vittad..
Material and methods
We synthesized Tuber malenconii ectomycorrhizae using carpophores gathered
in Villafranca (Castellon province, Spain). Part of the material was preserved
in the herbarium of the University of Alcala (AH-20004). We synthesized
ectomycorrhizae of T’ malenconii, T. aestivum, and T: mesentericum with
Quercus ilex subsp. ballota (Desf.) Samp. plants according to Bencivenga (1982)
and modified by Manjon & Garcia-Montero (1996). Plants were kept under
controlled environmental conditions in the Juan Carlos I Royal Botanical Garden
at the University of Alcala (Madrid, Spain). The degree of mycorrhization of each
plant was expressed as the number of Tuber mycorrhizae per total number of
apices counted (Bencivenga et al. 1987). Ectomycorrhizae were identified with
a stereoscopic microscope (photo-Leica WildMZ8) and compound microscope
(photo-Leica Leitz DMRB) following the recommendations proposed by Agerer
(1987-2002), Zambonelli et al. (1993), and Granetti (1995). Each character was
measured by taking measurements until regularity was attained in the values
observed.
Ectomycorrhizae
Our techniques for inoculating and producing mycorrhized plants, as well
as the substrates used, gave good results for both Tuber malenconii and
T. melanosporum. The six seed Quercus ilex subsp. ballota plants mycorrhized
with Tuber malenconii showed a 50% mycorrhization average (standard
deviation = 14) of all root apices when other fungal ectomycorrhizae were not
Tuber malenconii mycorthizae ... 409
present, while the six seed plants mycorrhized with T’ melanosporum averaged
a 62% (standard deviation = 16) coverage, also without ectomycorrhizae of
other fungi. The T: malenconii ectomycorrhizae (Figs. 1-4) were concentrated
above all in the proximal and median part of the root system.
Tuber malenconii Donadini, Riousset, G. Riousset & G. Chev. FIG. 1
FORM AND MODE OF RAMIFICATION: from slightly club-shaped to almost
cylindrical, or with monopodial-pinnate ramifications, without ramifications in
very young forms; UNRAMIFIED ENDS: 1500-3500 um long, 580-720 um diam,
straight, color ochre, darker in the central mycorrhiza, very clear in the apical
part, darkening in age; STRUCTURE OF THE SURFACE: smooth, with numerous
long cystidia that give it a woolly appearance, more abundant at the apex of the
young mycorrhizae and at the centre and base of the mature mycorrhizae; the
old mycorrhizae may be free of mycelium; RHIZOMORPHsS: absent.
Emanating hyphae
CYSTIDIATE with a very long sinuous shape, septate, unbranched; widths 2-3
um diam, lengths 15-30 um, very variable; colour yellowish. SURFACE smooth
or occasionally with mammellate incrustations, the termination is rounded and
simple but may be widened by the presence of these incrustations. HYPHAE
RAMIFICATIONS: the hyphal insertion point into the mantle may be simple,
slightly widened, or ramified. A single hypha may grow out of the mantle or
immediately branch into two forming a typical “Y” shape; sometimes two
hyphae emerge from one point. Such ramifications were observed only near
the hyphal insertion into the mantle, after which no further branching was
observed. PERCENTAGE OF THE KIND OF HYPHAE: on the basis of the counts
made we can establish 60% hyphae with non basal-forked ramifications (=
forked ramifications at the base), 30% cystidia with basal-forked ramifications,
and 10% doubtful hyphae due to difficulty of observation.
Anatomical characters of the outer mantle
STRUCTURE: pseudoparenchymatous with most of the pseudocells polygonal
with 4 to 5 sides, alternating with some cells with a sinuous form; DIMENSIONS
OF HYPHAL PSEUDOCELLS: (3)4-10(11) x (3)4-8(9) um.
Anatomical characters — cross section
THICKNESS AND DIFFERENTIATION: mantle is 10-30 um thick, a thin (3-
10 um thick) outermost layer can be differentiated where the hyphal cells are
more compact; CELL SHAPE AND DIMENSIONS: the outermost layer cells are very
difficult to measure; the remaining mantle is composed of cells that are either
rounded and then 8-15 x 4-9 um or angular with a tangential orientation.
A410 ... Garcia-Montero & al.
Tuber malenconii mycorrhizae ... 411
TANNIN CELLS 1-2 rows, sometimes forming a 3-6 um thick continuous
layer, individual cells averaging 5-14 x 2-5 tum. CoRTICAL CELLS of two shapes,
either rounded (18-24 x 18-22 um) or rectangular (35-50 x 20-30 um).
HARTIG NET present in 2-3 rows to a depth of 40-60 um; THICKNEss: 2-4 um;
SHAPE OF HYPHAL CELLS AND DIMENSIONS: rounded/rectangular, 2-4 x 1-1.5
Lum.
Table 1. Discriminating features among Tuber malenconii, T. aestivum, and
T. mesentericum ectomycorrhizae
SPECIES T. malenconii T: aestivum T: mesentericum
UNRAMIFIED ENDS
Length < 3500 um < 8500 um** < 3800 um**
Diameter < 720 um < 400 um** < 300 um**
MANTLE Polygonal pseudocells — Polygonal Polygonal pseudocells with
OUTER with rounded angles pseudocells with sharp angles*
STRUCTURE rounded angles* (secondarily diagnostic**)
CYSTIDIA
Form Without intercalary With intercalary Without intercalary
thickenings thickenings.* thickenings*
(sporadic**)
% with 30 % None** 5-10 % **
basal-forked
ramifications
* Zambonelli et al. (1993); ** Granetti (1995).
Discussion
Tuber malenconii ascocarps are readily diagnosed by their characteristic 8-
spored asci and ascospores with a regular reticulate-alveolate ornamentation
consisting of small, dense meshes. Unfortunately, the mycorrhizae of
T. malenconii, T. aestivum, and T: mesentericum are morphologically very
similar: all three produce numerous long cystidia that cover most of the
mycorrhizal surface and outer mantle pseudocells with a polygonal form
(Table 1; Zambonelli et al. 1993, Granetti 1995). In fact, T: aestivum and T:
mesentericum mycorrhizae are so similar (Fig. 2) that 7’ mesentericum can be
Fig 1 (left). Tuber malenconii (bar = 10 um). Fig. 1.1. Hyphal pseudocells from the outer surface
of the mantle with polygonal shape, with roundish angles (1000 x). Fig. 1.2. Hartig net (1000
x). Fig. 1.3. Macroscopic aspect of the mycorrhizae (100 x). Fig. 1.4. Detail of cystidium with
basal-forked ramifications (1000 x).
A12. ... Garcia-Montero & al.
Tuber malenconii mycorrhizae ... 413
separated from T! aestivum only by noting that a consistantly small percentage
(5-10%) of the cystidia fork at the bases (Di Massimo et al. 1995, Granetti 1995),
a character not seen in T: aestivum mycorrhizae. This forking is not confined to
T. mesentericum mycorrhizae, however, as an even greater percentage (30%) of
cystidia in T. malenconii mycorrhizae produce basal-forked ramifications (Fig
1.4). The differences in the degree of basal forking among three species suggest
that the character may have diagnostic potential in developing taxonomic keys
to all Tuber mycorrhizae. Such a key would be helpful in creating new quality
control protocols established to ensure that the mycorrhized plants have
not become contaminated by undesirable species during the truffle culture
process.
Acknowledgments
The authors are grateful to Drs. M. Castellano, P. Bonfante and A. Vizzini for serving as
pre-submission reviewers and for their valuable comments and suggestions. We thank
Margarita, Luis, Miriam, Pablo and P. Diaz for their support and collaboration. We
want to express also our gratitude to M* Cruz Gémez-Llano, Faustino Correas and the
personal of the University Library of E.T.S.I. Montes (UPM), and Prudence Brooke-
Turner for her linguistic assistance. This work have been sponsored by the INIA Project
of (SC94-129), by the DGICYT Project of (PB91-0165) and by the Fundacion Caja de
Madrid.
Literature cited
Agerer R. 1987-2002. Colour atlas of ectomycorrhizae. Einhorn-Verlag, Schwabsh-Gmund. 168
PP:
Alvarez IF, Parladé J, Trappe JM, Castellano MA. 1993. Hypogeous mycorrhizal fungi of Spain.
Mycotaxon 47: 201-217.
Bencivenga M, Ferrara AM, Fontana A, Granetti B, Gregori G, Lo Bue G, Palenzona M, Rebaudengo
E, Tocci A, Zambonelli A. 1987. Valutazione dello stato di micorrizazione di piante tartufigene.
Proposta di un metodo. Ministero dell’ Agricultura e Foreste, Roma. 14 pp.
Bencivenga M. 1982. Alcune metodiche di micorrizazione di piante forestali con il tartufo nero
pregiato di Norcia o di Spoleto (Tuber melanosporum Vittad.). L Informatore Agrario 38:
21155-21163.
Comandini O, Pacioni G. 1997. Mycorrhizae of Asian black truffles, Tuber himalayense and T.
indicum. Mycotaxon 63: 77-86.
Di Massimo G, Garcia-Montero LG, Bencivenga M, Manjon JL. 1996. Tuber indicum Cooke et
Massee, un tartufo orientale simile a T. melanosporum Vitt. Micol. Veget. Medit. 11: 107-114.
Di Massimo G, Garcia-Montero LG,. Manjon JL, Bencivenga M. 1995. Un nuovo carattere di
interesse diagnostico sulle micorrize di Tuber mesentericum. In H° Convegno Taxa e cenosi
fungine nell’ Area del Mediterraneo. Abstracts, 21 to 24 September 1995, Opi-Aquila, Italy.
Fig. 2. Tuber mesentericum (bars = 10 um). Fig. 2.1. Detail of cystidium with basal-forked
ramifications (1000 x). Fig. 2.2. Hyphal pseudocells from the outer surface of the mantle with
polygonal shape, with sharp angles (1000 x). Fig. 2.3. T. aestivum. Macroscopic aspect of the
mycorrhizae (100 x). Fig. 2.4. Hyphal pseudocells from the outer surface of the mantle with
polygonal shape, with roundish angles (1000 x) (the white segments represent 10 um).
414 ... Garcia-Montero & al.
Donadini JC, Riousset L, Riousset G, Chevalier G. 1979. Tuber malenconi nov. sp. Bull. Soc. Mycol.
France 94: 351-358.
Garcia-Montero LG, Di Massimo G, Manjon JL. 1996. Synthesis and description of Tuber malenconii
Don., Riouss. & Chev. ectomycorrhizae. pp. 51-52, in First International Conference on
Mycorrhizae. ICOM I. Abstracts, 4 to 9 August 1996, Berkeley, USA.
Garcia-Montero LG, Di Massimo G, Manjon JL, Garcia- Abril A. 2008. New data on ectomycorrhizae
of the Chinese truffles Tuber pseudoexcavatum and T. indicum. Submitted.
Granetti B. 1995. Caratteristiche morfologiche, biometriche e strutturali delle micorrize di Tuber di
interesse economico. Micol. Ital. 2: 101-117.
Granetti B. 1995. Caratteristiche morfologiche, biometriche e strutturali delle micorrize di Tuber di
interesse economico. Micol. Ital. 2: 101-117.
Manjon JL, Garcia-Montero LG. 1996. Sintesis micorricica con esporas de Tuber malenconii y otros
hongos con aplicacion forestal y en la truficultura. Bol. Soc. Micol. Madrid 21: 397-400.
Manjon JL, Garcia-Montero LG, Di Massimo G. 1998. Mycorrhizal synthesis of Tuber
pseudohimalayense and T. pseudoexcavatum, two truffles from Asia. p. 113, in Second
International Conference on Mycorrhizae. ICOM HU, Abstracts, 5 to 10 July 1998, Uppsala,
Sweden.
Montecchi A, Sarasini M. 2000. Funghi ipogei d’Europa. Associazione Micologica Bresadola,
Trento. 714 pp.
Riousset L, Riousset G, Chevalier G, Bardet MC. 2001. Truffes d'Europe et de Chine. Institut —
National de la Recherche Agronomique INRA, Paris. 181 pp.
Zambonelli A, Salomoni S, Pisi AM. 1993. Caratterizzazione anatomo-morfologica delle micorrize
di Tuber spp. su Quercus pubescens Willd. Micol. Ital. 22: 73-90.
Zambonelli A, Tibiletti E, Pisi AN. 1997. Caratterizzazione anatomo-morfologica della micorriza di
Tuber indicum Cooke & Massee su Pinus pinea L. and Quercus cerris L. Micol. Ital. 1: 29-36.
MYCOTAXON
Volume 105, pp. 415-419 July-September 2008
An albino form of Auricularia fuscosuccinea from
Lacandonia tropical forest, Chiapas, Mexico
SIGFRIDO SIERRA’, JOAQUIN CIFUENTES’,
FELIPE RUAN-SOTO? & RAMON MARIACA4
ssg@fciencias.unam.mx
'Lab. de Heterobasidiomycetes Tremeloides
*Lab. de Biodiversidad y Biogeografia Ecologica de Hongos
Fac. de Ciencias, UNAM. A.P. 70-181, Coyoacan, C.P. 04510 México, D.F
*Seccion de Micologia, Herbario Eizi Matuda
Escuela de Biologia, Universidad de Ciencias y Artes de Chiapas
* El Colegio de la Frontera Sur (ECOSUR)
San Cristobal de las Casas. Chiapas, México
Abstract - An albino form of Auricularia fuscosuccinea is described from Chiapas
tropical forest in Mexico. The taxonomy of Auricularia species is briefly reviewed.
Key words - Auriculariaceae, biological diversity, Neotropics.
Introduction
According to Ainsworth & Bisby’s Dictionary of the Fungi (Kirk et al. 2001),
there are approximately 15 recognized species of Auricularia Bull. ex Juss.,
represented by 124 species names listed in the Index of Fungi. Proliferation of
most species names occurred prior to Lowy (1951) at a time when species were
delimited based primarily on basidiome size, shape and color and the length
of abhymenial hairs (Saccardo 1888, 1895; Barrett 1910; Burt 1921; Kobayasi
1942). Teixeira (1945) utilized the medulla, a central zone of compact, parallel
hyphae that may be observed in transverse sections of basidiomata in addition
to the above features. Lowy (1951) suggested that the traditional characters of
basidiome size, shape and color varied with age and environmental conditions
and should, therefore, not be considered of primary taxonomic importance.
Instead, Lowy (1951, 1952) selected the presence/absence of the medulla, the
medullary width when present, and abhymenial hair length as features that
he believed to be stable and not significantly affected by age and changes in
environmental conditions. Lowy (1951, 1952) used these characters to delimit
species, recognizing ten species and reducing most of the names to synonymy.
416... Sierra & al.
While most workers (e.g. Cheng & Tu 1978, Kobayasi 1981) have followed
Lowy (1951, 1952), Olive (1958) and Raitviir (1971) questioned the value of
the characters utilized by Lowy (1951, 1952) to define species. |
Kobayasi (1981), who described 15 species and 5 forms of Auricularia, listed
three albino forms: A. auricula f. albicans (Berk.) Kobayasi (= A. auricula-judae
var. lactea Quél.), A. delicata f. alba Kobayasi, and A. polytricha f. leucochroma
(Kobayasi) Kobayasi. Li & Liu (1985) described one albino collection as a new
species, A. eburnea, and Wong (1989) described an albino form of A. cornea.
Albino forms of Auricularia have not been previously reported in neo-
tropical regions by Lowy (1971, 1980) nor in Mexico by Lowy (1965), Mendiola
& Guzman (1973), Pérez-Silva & Esqueda- Valle (1992) and Sierra & Cifuentes
(1993).
Material and methods
The specimens were collected and studied using standard mycological
techniques (Martin 1952, Lowy 1971, Cifuentes et al. 1986, Sierra 1992),
and with specimens deposited in FCME. Color numbers are from Methuen
Handbook of Colour (Kornerup & Wanscher 1978).
Taxonomy
Auricularia fuscosuccinea (Mont.) Henn.
“Lacandon albino form” FIGURE 1
Basidiomes orbicular, c. 8 cm in diameter, context thin. White when fresh
(Methuen 2A2 to white); semitransparent when dry. Upper surface smooth,
with scattered hairs 30-65 x 4-6 um, with acute tips. Medulla a single band
of hyphae centrally oriented. Basidia up to 60 x 3.5 um; basidiospores curved-
cylindrical, 10-13 x 4-5 um. Germination not observed.
EcoLoGy, RANGE AND DISTRIBUTION: Lignicolous, on unidentified rotten
wood. Found in hot and humid tropical forest habitat with rain year-round
except for a short dry season in March and April. Median annual precipitation
of 2400 mm.
SPECIMEN EXAMINED —~ MEXICO. CHIAPAS - Municipio de Ocosingo, Lacanja-
Chansayab, located at 16°46’08” north, 96°08'12” west, alt. 350 m — 24.1.2004 Ruan-Soto
46 (FCME 22561).
Figure 1: Auricularia fuscosuccinea. A. Basidiome (albino form). B. Basidiome (typical form).
C. Section of basidiome (abbreviations: ZP: Zona pilosa; ZC: Zona compacta; ZSS: Zona
subcompacta superioris; ZLS: Zona laxa superioris; M: Medullae; ZLI: Zona laxa inferioris;
ZSI: Zona subcompacta inferioris; H: Hymenium). D. Hairs. E. Medulla hyphae. F Hymenium.
G. & H. Spores (G. albino form). (H. typical form). Scale bar G & H=5 um.
A white form of Auricularia fuscosuccinea (Mexico) ... 417
418 ... Sierra & al.
ADDITIONAL SPECIMEN EXAMINED — MEXICO. CHIAPAS - Municipio de Ocosingo,
Lacanja-Chansyab. 18.VI.2004. Ruan-Soto 156 (FCME 22586), typical form of A.
fuscosuccinea.
CoMMENTS -— This albino form differs from the typical form only in colour.
A. fuscosuccinea “Lacandon albino form” is very easy to differentiate from the
albino form of A. cornea (= A. polytricha) described by Wong (1989) by its short
hairs (until 65 pm) and from A. auricula-judae v. lactea and A. delicata f. alba by
the presence of medulla. A. eburnea discribed by Li & Liu (1985) is different in
spores (15.6-18 x 5.5-7.5 um) and hair size (280-380(-450) um). This fungus
is rare. People in the region where it is found speak Lacandon (Levy-Tacher
2000). ‘The Lacandon name of this fungus is Sak’re lo’ro (Sak’re = white and lo’ro =
all species of Auricularia). The fungus is regarded as edible; the Lacandons eat it fried
in a “comal” (a kind of frypan) or prepare a dish with Auricularia and bean stock. This
is the first published record of a white Auricularia in Mexico.
Acknowledgements
We wish to thank Dr. Peter Roberts, Dr. George J. Wong and Dr. Peter Buchanan for
reviewing the manuscript and for helpful comments; Mr. Manuel Castellanos Chan
Kin for his helpful guidance in the rain forest. M.S. Diego Hernan Valencia Korosi for
reviewing the English manuscript. This research was supported by DGAPA IN-209605
and IN-218008.
Literature cited
Barrett ME 1910. Three common species of Auricularia. Mycologia 2: 12-18.
Burt EA. 1921. Some North American Tremellaceae, Dacryomycetaceae and Auriculariaceae. Ann.
Mo. Bot. Gard. 8: 361-396.
Cheng S, Tu CC. 1978. Auricularia spp. Pp. 605-625. In: Chang ST, Hayes WA. (Eds.) The biology
and cultivation of edible mushrooms. Academic Press, New York
Cifuentes J, Villegas M, Pérez-Ramirez L. 1986. Hongos. In: Lot A, Chiang F. (Eds.) Manual de
Herbario. Cons. Nac. Fl. Méx., A.C. México.
Kirk PM, Cannon PF, David JC, Stalpers JA. 2001. Ainsworth & Bisby’s Dictionary of the Fungi. 9th
ed, International Mycological Institute, CAB International, Wallingford.
Kobayasi Y. 1942. On the genus Auricularia from eastern Asia. Bull. Centr. Natl. Mus. Manchoukuo.
4: 19-35.
Kobayasi Y. 1981. The genus Auricularia. Bull. Natn. Sci. Mus. Tokio, Ser. B. 7(2): 41-67.
Kornerup A, Wanscher JH. 1978. Methuen Handbook of Colour. Eyre Methuen, Londres.
Levy-Tacher S. 2000. Sucesién causada por la roza - tumba — quema en las selvas de Lacanha,
Chiapas. Tesis de Doctorado. Colegio de Postgraduados. México.
Li L, Liu B, 1985. A new species of Auricularia. Journal of Shanxi University (Nat. Sci. Ed.) 1:56-
Do.
Lowy B. 1951. A morphological basis for classifying the species of Auricularia. Mycologia 43:
351-358.
A white form of Auricularia fuscosuccinea (Mexico) ... 419
Lowy B. 1952. The genus Auricularia. Mycologia 44: 656-692.
Lowy B. 1965. Estudio sobre algunos Tremellales de México. Bol. Soc. Bot. Mex. 29: 19-33.
Lowy B. 1971. Flora Neotropica, Monograph No. 6. Tremellales. Hafner. New York., 153 p.
Lowy B. 1980. Tremellales. Flora Neotropica No. 6 (Suppl.): 1-18.
Martin GW. 1952. Revision of the North Central Tremellales. Verlag Von J. Cramer, Lehre. 122 p.
Mendiola G, Guzman G. 1973. Las especies de Tremellales conocidas de México. Bol. Soc. Mex.
Mic. 7: 89-97.
Olive LS 1958. The lower Basidiomycetes of Tahiti - I. Bull. Torrey Bot. Club 85: 5-27.
Pérez-Silva E, Esqueda-Valle M. 1992. First record of jelly fungi (Dacrymycetaceae, Auriculariaceae
and Tremellaceae) from Sonora, México. Mycotaxon 44(2): 475-483.
Raitviir A. 1971. The tremellaceous fungi of the Far East. Pp. 84-154. In: Plants and animals of the
Soviet Far East. Valgus, Tallin.
Sacardo PA. 1888. Sylloge fungorum omnium hucusque cognitorum. Vol. 6. Padua, Italy. 928 p.
Sacardo PA. 1895. Sylloge fungorum omnium hucusque cognitorum. Vol. 11. Padua, Italy.753 p.
Sierra S. 1992. Contribucién al estudio taxondmico de algunos hongos tremeloides
(Heterobasidiomycetes) del centro y sur de México. Tesis de Licenciatura, Fac. de Ciencias,
UNAM. México, DE.
Sierra S, Cifuentes J. 1993. Contribucién al estudio taxondmico de los hongos tremeloides
(Heterobasidiomycetes) de México. Rev. Mex. Mic. 9: 119-137.
Teixeira R. 1945. Himenomicetos Brasileiros I. Auriculariales e Dacryomycetales. Bragantia 5:
154-186.
Wong GJ. 1989. Compatibility and fruiting studies of an albino form of Auricularia cornea.
Mycotaxon, 34(1): 259-266.
+ '
~
Volume 105, pp. 421-422 July-September 2008
Validation of Repetobasidiopsis and Trimitiella
(Basidiomycetes)
G.S. DHINGRA & AVNEET P. SINGH
dhingragurpaul@rediffmail.com
Department of Botany, Punjabi University
Patiala - 147 002, India
Abstract — In their original publications, Repetobasidiopsis grandisporus and Trimitiella
indica were referenced to holotypes deposited in two different herbaria. To ensure the
validity of the generic names Repetobasidiopsis and Trimitiella and the species names,
one herbarium (PAN) is designated for each holotype in accordance with ICBN Article
SS Be
Key words — nomenclature
Dhingra (2006) and Dhingra & Singh (2006) proposed the new genera
Trimitiella and Repetobasidiopsis to accommodate two new species, Trimitiella
indica and Repetobasidiopsis grandisporus respectively. In the protologues, two
different herbaria were cited for each holotype — PAN and LY for T! indica
and PAN and GH for R. grandisporus. However, Art. 37.7 of the International
Code of Botanical Nomenclature (McNeill et al. 2006) states, “For the name
of a new species or infraspecific taxon published on or after 1 January 1990 of
which the type is a specimen or unpublished illustration, the single herbarium
or collection or institution in which the type is conserved must be specified.”
Because two herbaria were designated for holotype collections for each type
species, the generic and species names were invalidly published.
Therefore we _ validate below Repetobasidiopsis, Repetobasidiopsis
grandisporus, Trimitiella and Trimitiella indica by designating the specimens
held in PAN as holotypes and those held in LY and GH as isotypes. According
to Art. 36.1, it is unnecessary to repeat the Latin diagnoses, but refer to those
published in Dhingra (2006) and Dhingra & Singh (2006).
Repetobasidiopsis Dhingra & Avneet P. Singh, gen. nov.
MycosBaNnk MB501457
“Repetobasidiopsis” Dhingra & Avneet P. Singh,
Mycotaxon 97: 115, 2006 (inval. Art. 37.7).
Typus: Repetobasidiopsis grandisporus Dhingra & Avneet P. Singh
422 ... Dhingra & Singh
Repetobasidiopsis grandisporus Dhingra & Avneet P. Singh, sp. nov.
MycosBank No. MB501362
“Repetobasidiopsis grandisporus” Dhingra & Avneet P. Singh,
Mycotaxon 97: 116, 2006 (inval. Art. 37.7).
Holotypus: G.S. Dhingra 19706 (PAN; IsoTyPeE at GH)
Trimitiella Dhingra, gen. nov.
MycoBaNnkK No. MB501458
“Trimitiella” Dhingra, Mycotaxon 97: 125, 2006 (inval. Art. 37.7).
Typus: Trimitiella indica Dhingra
Trimitiella indica Dhingra, sp. nov.
MycoBaNnk No. MB501367
“Trimitiella indica” Dhingra, Mycotaxon 97: 126, 2006 (inval. Art. 37.7).
Holotypus: G.S. Dhingra 19722 (PAN; IsoryPE at LY).
Acknowledgments
We thank Paul Kirk, Lorelei Norvell and Shaun Pennycook for alerting us to the need for -
validation and for providing reviews of this paper, and Head of Department of Botany,
Punjabi University Patiala for providing infrastructure.
Literature cited
Dhingra GS. 2006. Trimitiella gen. nov. (Basidiomycetes) from Eastern Himalaya, India. Mycotaxon
97: 125=128:
Dhingra GS, Singh AP. 2006. Repetobasidiopsis gen. nov. (Basidiomycetes) from Eastern Himalaya,
India. Mycotaxon 97: 115-118.
McNeill J, Barrie FE Burdet HM, Demoulin V, Hawksworth DL, Marhold K, Nicolson DH, Prado
J, Silva PC, Skog JE, Wiersema J, Turland NJ (eds.). 2006. International Code of Botanical
Nomenclature (Vienna Code). Adopted by the Seventeenth International Botanical Congress
Vienna, Austria, July 2005. Reg. Veget.146: i-xvi, 1-568.
MYCOTAXON
Volume 105, pp. 423-431 July-September 2008
New and unusual hyphomycetes
from Mahabaleshwar, India
J. PRATIBHA * & D.J. BHAT**
*jalmipratibha@rediffmail.com & **bhatdj@rediffmail.com
Department of Botany, Goa University
Goa-403 206, India
Abstract - Three conidial fungi, Tetraploa circinata sp. nov., Vamsapriya
mahabaleshwarensis sp. nov. and Gliophragma setosum, isolated from plant litter and
collected at Mahabaleshwar, Maharashtra, India, are described and illustrated.
Key words — biodiversity, anamorphic fungi, taxonomy, Western Ghat forests
Introduction
During the course of our studies on microfungi of the forests of Western
Ghats, two new dematiaceous hyphomycetes, Tetraploa circinata and
Vamsapriya mahabaleshwarensis, were isolated from fallen litter collected from
Mahabaleshwar, southern India. Tetraploa circinata is unique in its long, spirally
coiled, verrucose appendages. Vamsapriya mahabaleshwarensis differs from the
only previously described species in this genus by its branched conidial chains
developing on polytretic conidiogenous cells. Gliophragma setosum, a rare
conidial fungus, was also isolated during the course of this study.
Taxonomic descriptions
Tetraploa circinata Pratibha & Bhat, sp. nov. Figs. 1, 2
MycoBank MB 512004
Ad fungos conidiales, hyphomycetes. Coloniae in substrato naturali effusae, atrobrunneae
vel nigrae. Mycelium superficialia, ex hyphis laevibus, pallide brunneis, ramosis, septatis,
2.5-3 ym lat. compositum. Setae et hyphopodia absentia. Conidiophora micronematica,
ramosa, atrobrunnea. Cellulae conidiogenae monoblasticae, incorporatae. Conidia ex
corpore muriforme et appendicibus quatuor composita, solitaria, sicca, atrobrunnea,
conidiorum sunt 2 formae; conidia priora ex 4 columnis compositum, 13-15 x 21-26
*Current address: Myko Tech Pvt. Ltd. [Plot No. 12, Mapusa Industrial Estate,
Dattawadi Mapusa, Goa 403507, India]
APA ... Pratibha & Bhat
Fic. 1. Tetraploa circinata. Two types of conidia.
um, columnae 5-7 yum lat., divergentibus ad apicem, appendices spirales, verruculosae,
atrobrunneae, 54-415 x 3-3.5 um; conidia cetera ex 4 columnis compositum, 15-24 x
12-22 um, appendices laeviae, 5-22 um longa, 2-4 ym lat.; columnae 8-10 ym lat.
Ho otyPe: On dead, decaying bamboo twig, 04/11/2006, J. Pratibha, Mahabaleshwar,
Maharashtra, India, Herb. No. HCIO 48133 (Isotype: GUBH MH3)
New hyphomycetes from Mahabaleshwar (India) ... 425
Fic. 2. Tetraploa circinata.
a-d. First type of conidia with long, spiral, verrucose, appendages;
e-g. Second type of conidia with short, smooth appendages.
426 ... Pratibha & Bhat
Conidial fungi, hyphomycetes. Colonies on natural substrate effuse, dark
brown to black: Mycelium partly superficial, composed of septate, branched,
light brown, smooth hyphae 2.5-3 um wide. Setae and hyphopodia
absent. Conidiophores micronematous, branched, dark brown, forming
an entanglement on the substrate. Conidiogenous cells monoblastic,
undifferentiated from conidiophores. Conidia consisting of a dictyosporous
body and four appendages, solitary, dry, dark brown, of two morphological
types; first type 13-15 um long, 21-26 um wide, consisting of four columns each
of 1-2 cells, columns 5-7 um wide, each terminating apically in a long, spiral
or sinuate, verrucose, dark brown appendage, 54-415 x 3-3.5 tum; second type
of conidia 15-24 um long, 12-22 um wide, consisting of four columns each of
2-5 cells, columns 8-10 um wide, with short, smooth, apical appendages 5-22
um long and 2-4 um wide.
Notes: The genus Tetraploa was established by Berkeley & Broome in 1850 with
T. aristata Berk. & Broome as type species. So far 11 species and 2 varieties are
accommodated in the genus: T: abortiva, T. aristata var. aristata, T. aristata var.
sacchari, T. biformis, T. curviappendiculata, T. divergens, T. ellisii, T. javanica, -
T. longissima, T. muscicola, T. scabra and T. setifera (Revay 1993, Matsushima
& Matsushima 1996, Hatakeyama et al. 2005, Anonymous 2008). The genus
is characterized by conidia that are sessile, solitary, brown, verruculose, tetra-
columnar, with shallow furrows between the columns; the columns diverge
from one another apically and each terminates in an arm or appendage which
is setiform, septate and tapering at the tip (Ellis 1949, 1971). Of the known
species, T. aristata, with two types of conidia and long appendages, bears some
similarity to T. circinata. In T. aristata, the first type of conidia are 25-39 x
14-29 um with smooth appendages 12-80 x 4.5-8 um while the second type
of conidia, which consist of only 2 columns of cells, are 8-18 x 7-12 um with
smooth appendages 80-330 x 3-6 um. In T. circinata the first type of conidia
are 13-15 x 21-26 um with long, spiral, verrucose appendages, 54-415 x 3-3.5
um and the second type of conidia are 15-24 x 12-22 um with short, smooth
appendages, 5-22 x 2-4 um.
Vamsapriya mahabaleshwarensis Pratibha & Bhat, sp. nov. Figs. 3, 4
MycoBank MB 512005
Ad fungos conidiales, hyphomycetes. Coloniae in substrato naturali effusae, atrobrunneae
vel nigrae. Mycelium substrato immersum, ex hyphis laevibus, pallide brunneis, ramosis,
septatis, 2.5-3 um latis compositum. Setae et hyphopodia absentia. Synnematica erecta,
recta vel flexuosa, atrobrunneae vel nigrae, apice divergentibus composita, 655-1150 um
longa, 100-150 um lat. ad basim, 15-45 um lat. ad medio, 30-80 um lat. ad apicem.
Conidiophora macronematica, synnematica, septata, ramosa, atrobrunneae, laevia.
Cellulae conidiogenae polytreticae, integratae vel discretae, terminaliae vel intercalares,
New hyphomycetes from Mahabaleshwar (India) ... 427
Fic. 3. Vamsapriya mahabaleshwarensis. Synnemata, conidiogenous cells and conidia.
nunquam cicatricem, pallide brunneae, 6-23 x 3-5 um. Conidia sicca, catenulata, laevia
vel verruculosa, 0-4-septata, ad septa constricta, ovoidea, ellipsoidea vel oblonga, utrinque
rotundata, 5-25 x 4-9 um.
HoLortype: On unidentified dead twig, 04/11/2006, J. Pratibha, Mahabaleshwar,
Maharashtra, India, Herb. No. HCIO 48134. (Isotype: GUBH MH4)
Conidial fungi, hyphomycetes. Colonies on natural substrate effuse, dark
brown to black. Mycelium immersed in the substrate, composed of septate,
branched, light brown, smooth, 2.5-3 um wide hyphae. Setae and hyphopodia
absent. Synnemata erect, straight to flexuous, dark brown to black, diverging
at the apex, 655-1150 um long, 100-150 um wide at the base, 15-45 um wide
in the middle, 30-80 um wide at the apex. Conidiophores macronematous,
A28 ... Pratibha & Bhat
Fic. 4. Vamsapriya mahabaleshwarensis.
a. Synnemata; b-c. Fertile apical region;
d-f. Conidiophores with polytretic conidiogenous cells and conidia;
g-j. Catenate conidia with branched chains.
New hyphomycetes from Mahabaleshwar (India) ... 429
synnematous, septate, branched, dark brown, smooth. Conidiogenous cells
polytretic, integrated or discrete, terminal or intercalary, non-cicatrized, light
brown, 6-23 x 3-5 um. Conidia dry, catenate, smooth to minutely verruculose,
0-4-septate, constricted at the septa, ovoid, ellipsoidal or oblong, rounded at
both ends, 5-25 x 4-9 uum, developing in branched chains which secede easily.
Notes: The monotypic genus Vamsapriya was established by Gawas & Bhat, with
V. indica as type, to accommodate conidial fungi with catenate, phragmosporous
conidia on synnematous conidiophores with non-cicatrized, monotretic,
conidiogenous cells (Gawas & Bhat 2005). Vamsapriya mahabaleshwarensis
differs from the type species by formation of polytretic conidiogenous cells
and branched conidial chains. Conidia in V. indica are smooth, 2—12-septate,
10-80 x 4-6 um, whereas in V. mahabaleshwarensis they are smooth to minutely
verruculose, mostly 0-4-septate, rarely up to 10-septate, 5-25 x 4-9 um.
Gliophragma setosum Subram. & Lodha, Can. J. Bot. 42: 1060. 1964. Fig. 5
Conidial fungi, hyphomycetes. Colonies on natural substrate effuse, cottony,
grayish, velvety. Mycelium party superficial on the substrate, composed of
septate, branched, hyaline, smooth hyphae 2-2.5 um wide. Synnemata erect,
straight to flexuous, brown, 105-150 x 12-25 um, enclosing the lower part
of setae. Setae branched, septate, smooth, light brown, 230-270 x 2.5-4 um;
middle seta unbranched, dark brown, thick-walled, 260-300 x 3-6.5 um.
Conidiophores macronematous, synnematous, septate, branched, light brown,
smooth, 170-250 x 3-5 um, developing together with setae. Conidiogenous
cells polyblastic, integrated. Conidia aggregated in slimy, light brown masses;
individual conidium hyaline, smooth, consisting of 5-7 cells separated by
narrow isthmi, 40-48.5 um long, 3-4.5 um wide.
SPECIMEN EXAMINED: On leaf litter, 04/11/2006, J. Pratibha, Mahabaleshwar,
Maharashtra, India, Herb. No. HCIO 48135; GUBH MH12.
Notes: Gliophragma setosum originally isolated from horse dung from
Massoorie, Uttar Pradesh, India, is the type and only species of the genus
Gliophragma Subram. & Lodha (Subramanian & Lodha 1964). We isolated the
fungus from plant litter collected at Mahableshwar, India, and this is only the
second known collection.
Acknowledgments
We thank the University Grants Commission, Council of Scientific and Industrial
Research and Ministry of Environment & Forests, Government of India, New Delhi,
for financial support in the form of research grants during the tenure of which this
work was carried out. The authors thank Prof. Bryce Kendrick, Canada, and Dr. Eric
McKenzie, New Zealand, for reviewing the manuscript.
A430 ... Pratibha & Bhat
Fic. 5. Gliophragma setosum.
a. Setose synnema; b-d. Conidiogenous cells with conidia;
e-f. Conidia with isthmi.
‘op
New hyphomycetes from Mahabaleshwar (India) ... 431
Literature cited
Anonymous. 2008. CABI Bioscience, CBS, LandcareResearch, Index Fungorum. Retrieved 16 June
2008, from http://www.indexfungorum.org/Names/Names.asp
Ellis MB. 1949. Tetraploa. Transactions of the British Mycological Society 32: 246-251.
Ellis MB. 1971. Dematiaceous Hyphomycetes. Commonwealth Mycological Institute, Kew, Surrey,
England.
Gawas P, Bhat DJ. 2005. Vamsapriya indica gen. et sp. nov., a bambusicolous, synnematous fungus
from India. Mycotaxon 94: 149-154.
Hatakeyama S, Tanaka K, Harada Y. 2005. Bambusicolous fungi in Japan (5): three species of
Tetraploa. Mycoscience 46: 196-200.
Matsushima K, Matsushima T. 1996. Fragmenta mycologica - II. In: Matsushima T (ed) Matsushima
mycological memoirs, no. 9. Kobe, Japan, pp. 31-40.
Revay A. 1993. Some new or interesting hyphomycetes from Hungary. Nova Hedwigia 56:
473-482.
Subramanian CV, Lodha BC. 1964. Two interesting Hyphomycetes. Canadian Journal of Botany
42: 1057-1063.
MYCOTAXON
Volume 105, pp. 433-439 July-September 2008
Record of the rare greenhouse fungus Lepiota elaiophylla
(Agaricales, Agaricaceae) in Prague, Czech Republic,
with notes on its taxonomy and distribution
JAN HOLEC? & VACLAV HALEK?
jan_holec@nm.cz
‘National Museum, Mycological Department
Vaclavské nam. 68, Prague 1, CZ 115 79, Czech Republic
?Pomoranskd 482, Prague 8, CZ 181 00, Czech Republic
Abstract — A find in Prague (Praha) of Lepiota elaiophylla, a recently described species
growing in greenhouses and containing amanitins, is published. The fungus was found
on soil in a large flower pot with a horticultural variety of Guzmania sp. (Bromeliaceae),
bought in a flower shop in Prague and later kept in a heated room of a rectory. Macro-
and microcharacters are presented and discussed and the substrate and habitat are
described in detail. The world distribution is summarized. To date, L. elaiophylla is
known from 6 countries of western, central and southern Europe and from natural
habitats in the Neotropics of Brazil.
Key words — Basidiomycota, systematics, mycogeography, Central Europe
Introduction
Lepiota elaiophylla was described as a new species quite recently (Vellinga &
Huijser 1998). It was distinguished from its counterpart Lepiota xanthophylla
P.D. Orton by structure of the pileus covering, shape of the cheilocystidia and
colours of the basidiocarps (Vellinga & Huijser 1998, Vellinga 2001a). In the
protologue, collections from the Netherlands were cited. Moreover, several
records from Europe published as L. xanthophylla are considered to be in fact
L. elaiophylla (Vellinga & Huijser 1998). They originate from Germany (Besl et
al. 1984), Italy (Migliozzi et al. 1989) and Switzerland (Breitenbach & Kranzlin
1995: no. 236). Recently, L. elaiophylla was found in Austria, in the tropical
greenhouse of the Botanical Garden in Graz and in a glasshouse of the Botanical
Garden in Vienna (Pidlich-Aigner & Hausknecht 2001, Pidlich-Aigner et
al. 2002) and in a tropical greenhouse of a butterfly farm in Mecklenburg-
Vorpommern, Germany (Kreisel 2007). In all these countries, L. elaiophylla
was found exclusively in greenhouses or hothouses whereas L. xanthophylla
434 ... Holec & Halek
grows in deciduous forests in temperate Europe (Vellinga & Huijser 1998).
Concerning the occurrence of L. elaiophylla in the wild, Vellinga & Huijser
(1998) discuss a record by Pegler (1977) from east Africa; however, there are
some discrepancies in several microcharacters. Records from natural habitats
in Brazil were published by Meijer (2006: state of Parana) and Wartchow et al.
(2008: state of Pernambuco).
As shown by Besl et al. (1984), L. elaiophylla (misidentified as L. xanthophylla,
see Vellinga & Huijser 1998) contains a-amanitin and B-amanitin.
Phylogenetic analysis of Lepiota (Vellinga 2001b, 2003) showed that L.
elaiophylla belongs to a different clade than L. xanthophylla and that this clade
also harbours other amanitin containing species like L. subincarnata J.E. Lange
and L. brunneoincarnata Chodat & C. Martin.
The aim of this paper is to describe macro- and microcharacters of the
Prague record of L. elaiophylla and to summarize the recent information on its
taxonomy and distribution.
Material and methods
All microscopic examinations were made using an Olympus BH-2 microscope.
The observations and measurements were made on material mounted in a
5% KOH solution. Iodine reaction was studied in Melzer’s reagent prepared
according to the formula given in Moser (1983). For spore size measurements,
randomly selected mature spores were used. Spores were measured without
the hilar appendix (apiculus). The expression [41,2,1] means that 41 spores
from 2 basidiocarps originating from 1 collection were measured. Illustrations
of microcharacters were drawn at a magnification of 1250 x using a drawing
tube. For micromorphological terminology see Bas et al. (1988). The collection
studied is kept at the Mycological Department of the National Museum, Prague,
Czech Republic (herbarium PRM).
Results
Lepiota elaiophylla Vellinga & Huijser PLATE 1
Bollettino del Gruppo Micologico G. Bresadola —- Nuova Serie 40(2-3): 462, 1998 (“1997”).
MACROCHARACTERS (of slightly dried-up fruitbodies brought to National
Museum, Prague, by the second author) — pILEUs 5-23 mm, hemisphaerical
with involute margin when young, then convex, dry, brown tomentose
when young, then disrupting into densely arranged, adpressed, tomentose,
up to 1 mm broad ochre-brown, olive-brown to brown scales on lemon-
yellow background; LAMELLAE medium dense, L = 40-50, | = 1, ventricose,
lemon yellow, lamellar edge irregular, concolorous; stipE 20-45 x 2-4 mm,
cylindrical, hollow, without ring, with lemon-coloured arachnoid veil when
Lepiota elaiophylla in the Czech Republic ... 435
young, which disappears with age, upper quarter to third glabrous, pale lemon-
yellow, towards base with concolorous background but covered with densely
arranged scales having the same character as those on pileus surface; CONTEXT
pale lemon-yellow in pileus, whitish with lemon tinge in stipe; sMELL musty,
like a cellar; TASTE mild.
MICROCHARACTERS — BASIDIOSPORES (6.5)6.8-8.0(8.5) x (3.2)3.5-4.0
tum [41,2,1], mean value 7.2 x 3.6 um, Q = 1.70-2.22, average 2.00, in face
view oblong to subcylindrical, in side view with slight suprahilar depression,
with short hilar appendix, slightly thick-walled, smooth, hyaline in water or
KOH, distinctly dextrinoid in Melzer’s reagent (wall yellow-brown, interior
pale brown-violet); BAsIDIA 22-32 x 7-8 um (without sterigmata), clavate,
4-spored; BASIDIOLAE 17-25 x 5-8 um, clavate; LAMELLAE EDGE almost
sterile, rarely with basidia; CHEILOCYSTIDIA numerous, 1- to 3-celled (with
0-2 basal cells which are short and cylindrical), terminal cells 17-30 x 5-10
um, narrowly clavate to clavate, thin-walled, hyaline; PLEUROCYSTIDIA absent;
LAMELLAR TRAMA regular to subregular, composed of parallel to subparallel
hyphae 2-10(12) um broad, cells cylindrical and long or slightly inflated and
short (15-40 um), hyaline, with yellow membranal pigment, subhymenium
made up of densely arranged, short, branched, septate cells; PILEUS CUTICLE
a thick intricate trichoderm, composed of long, interwoven, flexuous hyphae
mostly composed of 1-3 cells, basal cells short, cylindrical or inflated, flexuous,
terminal cells long, 50-150 x 9-13 um, flexuose, cylindrical, narrowly fusiform
or narrowly clavate, slightly thick-walled, with up to 0.5 um thick wall, with
homogeneous yellow membranal pigmentation; PARTIAL VEIL (looking like
cortina between pileus margin and upper part of stipe) composed of parallel
to slightly interwoven hyphae, 3-6 um broad, cylindrical, thin-walled, finely
yellow-brown incrusted; sCALEs on stipe surface composed of cells resembling
those on pileus surface, terminal cells long, interwoven, flexuous, cylindrical or
narrowly clavate, slightly thick-walled, with up to 0.5 um thick wall, basal cells
shorter and narrower, cylindrical; CLAMP CONNECTIONS present in all tissues.
Hasitat — L. elaiophylla was found on soil in a large flower pot with a
horticultural variety of Guzmania sp. (Bromeliaceae) in the year 2007. The
flower pot with the plant was bought in about 1997 in a flower shop in Prague,
Lazarska street. The origin of the plant and of its substrate (soil bought in a
flower shop) is unknown. In about 2003, the Guzmania was replanted into a
larger flower pot with garden substrate of unknown origin. Till 2006, the flower
pot was kept in a private house in a room where it was hot in summer (25-35
°C) and without heating in winter (average temperatures about 15 °C). During
this period the fungus was not observed. In 2007, the flower pot was moved to
the rectory in Prague-HoleSovice and kept in a heated room with rather stable
436 ... Holec & Halek
Lettre egant
U
Plate 1. Microcharacters of Lepiota elaiophylla (PRM 858090).
B: basidiospores, CH: cheilocystidia, PC: pileus cuticle. Bar = 10 pm.
average temperature (about 23 °C). However, the fructification of L. elaiophylla
was observed on hot August days when the temperature was higher (up to 30
°C). During the whole period 1997-2007, Guzmania was normally watered (no
special treatment).
MATERIAL STUDIED — CZECH REPUBLIC. PRAGUE (PRAHA), city part Prague-
Holegovice, street Dukelskych hrdint 54, RECTORY OF THE CHURCH OF Sv. ANTONIN (ST.
ANTHONY), on soil in large flower-pot inside a room, 16.VIII.2007 leg. Matus Kocian,
det. J. Holec, rev. E. C. Vellinga (PRM 858090).
Lepiota elaiophylla in the Czech Republic ... 437
Discussion
Taxonomy
Basidiocarps of L. elaiophylla found in Prague agree in all important
characters with the original material from the Netherlands described by
Vellinga & Huijser (1998). The conspecificity was also confirmed by E.C.
Vellinga who obtained the material for revision. The pileus colour given in the
original description is more vivid (dark orange-brown or reddish-ochraceous
brown at centre). However, the colours of Prague basidiocarps were observed
in slightly dry state one day after the removal of the basidiocarps from the
flower-pot when the colours were not so bright anymore. Mean values of spore
size of Prague material are slightly smaller than in the original description
but they are within the range given by Vellinga & Huijser (1998). Except for
these small differences, all diagnostic characters of L. elaiophylla (characters
of the similar species L. xanthophylla P. D. Orton are given in brackets) are
present: absence of a distinct layer of clavate elements in pileus cuticle (present
in L. xanthophylla), clavate cheilocystidia (lageniform, utriform to clavate in
L. xanthophylla), slightly smaller spores (up to 10 x 5 um in L. xanthophylla),
olive to lemon-yellow tinge on pileus surface (yellow in L. xanthophylla) and
dirty olive-brown colour of dry lamellae (yellow in L. xanthophylla). The same
characters were also observed by Pidlich-Aigner & Hausknecht (2001).
Distribution and ecology
To date, Lepiota elaiophylla is known only from 6 countries of western,
central and southern Europe (The Netherlands, Germany, Switzerland, Austria,
Czech Republic, and Italy; for references see Introduction) and from Brazil
(Meijer 2006, Wartchow et al. 2008). L. elaiophylla is included in the red list
of Finnish fungi (Rassi et al. 2001) in the category of not evaluated taxa and
without any information on its record in this country. According to Hansen &
Knudsen (1992), the species is not known from Nordic countries, which was
confirmed by J. Vesterholt, the editor of a new edition of Nordic mycological
flora (pers. comm., 2008).
The identity of a potential record from east Africa (Pegler 1977, as
L. xanthophylla) is uncertain (Vellinga & Huijser 1998). The record by Saar
(2001) from Germany needs a revision (if a voucher specimen exists) as the
published photograph suggests that the fungus could be L. xanthophylla. All
European records originate from greenhouses, hothouses (often in botanical
gardens) or rooms and the fungus was found on horticultural substrate used
for cultivated plants. It is apparent that the fungus requires relatively high
temperature for its fructification. This agrees well with its records from natural
habitats in the Neotropics of Brazil. In the state of Parana, the fungus was found
in a mixed ombrophilous forest and dense ombrophilous forest at an altitude
438 ... Holec & Halek
up to 900 m (Meijer 2006), in the state of Pernambuco it was growing solitary
on soil among litter in a tropical forest (Wartchow et al. 2008). The species
appears to be widespread, as Pernambuco and Parana are quite far away from
each other.
Concerning the European records, it would be interesting to know whether
the fungus is distributed via mycelium in horticultural substrate of tropical or
subtropical origin (or in substrate of other tropical plants moved from nature to
the greenhouses) or via air transport of spores. At this moment it is impossible
to solve this question as in all published records the origin of its substrate is
unknown. It is also interesting to know how long the spores are viable, and how
long it takes for the mycelium to produce fruitbodies.
Acknowledgements
We would like to thank the reviewers of this paper, Else C. Vellinga (Berkeley, USA)
and Vladimir Antonin (Brno, Czech Republic), for their valuable comments. The
work was financially supported by the Ministry of Culture of the Czech Republic
(MK00002327201).
Literature cited
Bas C, Kuyper TW, Noordeloos ME, Vellinga EC (eds.). 1988. Flora agaricina neerlandica. Vol. 1.
A.A. Rotterdam/Brookfield, A.A. Balkema.
Besl H, Mack P, Schmid-Heckel H. 1984. Giftpilze in den Gattungen Galerina und Lepiota.
Zeitschrift fir Mykologie 50(2): 183-192.
Breitenbach J, Kranzlin F. 1995. Pilze der Schweiz. Vol. 4. Luzern, Verlag Mykologia.
Hansen L, Knudsen H. 1992. Nordic macromycetes. Vol. 2. Copenhagen, Nordsvamp.
Kreisel H. 2007. Beitrage zur Pilzflora von Mecklenburg-Vorpommern (Deutschland). Feddes
Repertorium 118(3-4): 113-128.
Meijer AAR de. 2006. Preliminary list of the macromycetes from the Brazilian state of Parana.
Boletim do Museu Botanico Municipal 68: 1-55.
Migliozzi V, Brunori A, Coccia M. 1989. La micoflora delle serre di S. Sisto Vecchio in Roma.
Lepiotee (2a parte). Rivista di Micologia 32(3-4): 99-111.
Moser M. 1983. Die Rohrlinge und Blatterpilze. 1-533, in Kleine Kryptogamenflora, vol. 2b/2, ed.
5. Jena, Gustav Fischer Verlag.
Pegler DN. 1977. A preliminary agaric flora of East Africa. 1-615, in Kew Bulletin Additional
Series, vol. 6. London, Royal Botanic Gardens Kew.
Pidlich-Aigner H, Hausknecht A. 2001. Grofspilze in den Gewachshausern des Botanischen
Gartens der Universitat Graz. Osterreichische Zeitschrift fiir Pilzkunde 10: 43-73.
Pidlich-Aigner H, Hausknecht A, Scheuer C. 2002. Annotated list of macromycetes found in the ©
greenhouses of the Botanic Garden of the Institute of Botany in Graz. (Austria), 1998-2001.
Fritschiana 32: 49-61.
Rassi P, Alanen A, Kanerva T, Mannerkoski I (eds.). 2001. The red list of Finnish species. Helsinki,
Ministry of the Environment & Finnish Environment Institute.
Saar G. 2001. Mykologische Beobachtungen in 3 Garten. Tintling 26: 52-63.
Vellinga EC. 2001a. 7. Lepiota (Pers.: Fr.) S.F. Gray. 109-151, in ME Noordeloos, TW Kuyper,
Lepiota elaiophylla in the Czech Republic ... 439
EC Vellinga (eds.), Flora agaricina Neerlandica, vol. 5. Lisse etc., A.A. Balkema Publishers.
Vellinga EC. 2001b. Studies in Lepiota III. Some species from California, U.S.A. Mycotaxon 80:
285-295.
Vellinga EC. 2003. Phylogeny of Lepiota (Agaricaceae) - evidence from nrITS and nrLSU sequences.
Mycological Progress 2(4): 305-322.
Vellinga EC, Huijser HA. 1998 (“1997”). Lepiota xanthophylla and its greenhouse counterpart.
Bollettino del Gruppo Micologico G. Bresadola — Nuova Serie 40(2-3): 457-464.
Wartchow E, Putzke J, Cavalcanti MAQ. 2008. Agaricaceae Fr. (Agaricales, Basidiomycota) from
areas of Atlantic forest in Pernambuco, Brazil. Acta Botanica Brasilica 22(1): 287-299.
MYCOTAXON
Volume 105, pp. 441-446 July-September 2008
Morchella tomentosa, a new species from
western North America, and notes on M. rufobrunnea
MICHAEL Kuo
mushroomexpert@yahoo.com
POT Boxwa7 12
Charleston, IL 61920 USA
Abstract — A new species of Morchella, M. tomentosa, is described from western North
America; the range of M. rufobrunnea, previously documented from Mexicos Gulf
Coast, is extended to California and Oregon; the history of taxonomic names applied to
M. rufobrunnea on the West Coast is discussed.
Key words — morels, taxonomy
Introduction
Over 500 North American collections of Morchella were solicited from amateur
collectors or made by the author from 2002 to 2007 as part of an effort known
as the Morel Data Collection Project (MDCP). Preliminary molecular results
(see Kuo 2005, 2006) revealed over a dozen genetically distinct morels among
the collections. While several of these putative species are morphologically
cryptic, as yet poorly documented, or otherwise insufficiently understood, two
morels from western North America are easily distinguished morphologically
and ecologically. One is M. rufobrunnea; the other, M. tomentosa, has been
informally described (Kuo 2005, 2006, McFarlane et al. 2005, Pilz et al. 2004,
2007) but not previously given formal taxonomic status.
Taxonomy
Morchella tomentosa M. Kuo, sp. nov. PLATE 1
MycoBAnk MB 511840
Ascomata (40)50-120 mm alta; capitulum subconicum vel subcylindricum vel ovoideum;
costae tomentosae, canae vel atrae, interdum pallescens; hymenium atrum, interdum
pallescens; stipes tomentosus, ater, interdum pallescens; pili 100-400 x 7-28 um; bruni
in 2% KOH.
Ho.totypus. Biotopium in silvis coniferibus incensis; 1,200 m. altitudinis; USA, in
Montanaense, ad Mineral County; S. Engstrom col.; MDCP 06150405; specimen typicum
in Herb. F. conservatum.
442 ... Kuo
AscoMATA (40)50-120 mm high. HyMENopPHORE (20)30-110 mm high;
(15)30-50 mm wide at the widest point; subconic to subcylindric or ovoid;
pitted and ridged; with 18-26 primary vertical ridges and numerous shorter,
secondary vertical ridges and transecting horizontal ridges; attached to stipe
with a sinus about 1-2 mm deep and 1-2 mm wide. RipGEs densely tomentose;
black, silvery black, brownish black, brownish gray, watery gray, brown,
or brownish when young; becoming gray, grayish, pale tan, or whitish with
maturity; bluntly rounded when young, but with age often flattened or eroded,
revealing the whitish context beneath. Pirs primarily vertically elongated by
maturity but irregularly shaped when young; glabrous; opening and deepening
with development; progressing from gray or nearly black when immature
to gray, watery gray, brown, yellowish tan, or nearly white (especially when
exposed to prolonged sunlight) at maturity. Srrp—E 20-60 mm high; 10-40
mm wide; often basally clavate to subclavate; flared slightly to apex; densely
tomentose and gray to black when young, becoming finely tomentose or nearly
glabrous and gray to tan, watery brownish, or nearly white, with brownish
tufts of separated tomentum. CONTEXT whitish; 1-5 mm thick in the hollow -
hymenophore; in the stipe chambered or layered. STERILE INNER SURFACE
whitish and pubescent.
CHEMICAL REACTIONS (tested on MDCP 08070404, two days after collection):
Ammonia very pale orange, then grayish on sterile inner surface and context,
negative on hymenium; KOH (5%) negative on all surfaces; FeSO4 negative on
all surfaces.
Ascospores elliptical; smooth; contents homogeneous; (15)18-20(22.5) x
(7.5)8-12.5(15) um; average Q=1.87. ASCI 8-spored; 225-290 x 12.5-20 um;
cylindric; hyaline. PARAPHYSES cylindric to subclavate; apices rounded to
subacute; with 2-5 septa; hyaline or with brown to brownish homogeneous
contents in KOH (2%); (125)150-175(225) x 5-10(15) um. RESIDUAL
PARAPHYSES On Sterile ridges similar to paraphyses; hyaline or with brownish
contents; bundled. HAIRS ON STERILE RIDGES projecting from residual
paraphyses; variable in shape (cylindric, clavate, subclavate, lageniform,
or subcapitate); 120-250(400) x 7.5-15(25) um; abundant; with brown to
brownish contents in KOH (2%); septate; often with thickened walls. HArrs ON
STIPE SURFACE abundant; cylindric to subclavate; septate; 100-275 x 10-17 um;
with brown walls in KOH (2%); contents hyaline to brownish.
EcoLocy — Appearing at altitudes of 1,000-3,400 m in lightly to moderately
burned conifer forests including forests dominated by Pseudotsuga menziesii
(Mirb.) Franco on the West Coast and forests dominated by Picea engelmannii
Parry and Abies lasiocarpa (Hook.) Nutt. or Pinus contorta var. latifolia Engelm.
in the Rocky Mountains. Found primarily in years following forest fires but
Morchella tomentosa sp. nov. (North America) ... 443
PiaTE 1. Morchella tomentosa. A: Pigmented hairs on stipe surface (scale: x 2.5 um). B: Tomentose
ridges of young hymenophore. C: Holotype specimen (about half life size). D, E, F: Isotype
specimens (about half life size), demonstrating morphological variability.
often appearing in dwindling numbers for several seasons thereafter; Alaska,
the Yukon Territories, Montana, Idaho, Oregon, and Colorado; May through
August.
SPECIMENS EXAMINED — UNITED STATES OF AMERICA: MONTANA—05300402
(May 30, 2004; Ravalli County, S. Schwartzman); 06100102 (June 10, 2001; Ravalli
County, J. Holmes); 06150402, 06150403, 06150405 HOLOTYPE, 06150406, 06150407,
06150409 (June 15, 2004; Mineral County, S. Engstrom); 06170402 (June 17, 2004;
Missoula County, S. Engstrom); 06180401, 06180402, 06180403, 06180404, 06180405,
06180406, 06180407, 06180408, 06180409, 06180413, 06180414, 06180415, 06180420,
06180421 (June 18, 2004; Mineral County, S. Engstrom); 06300401, 06300402, 06300403,
06300404, 06300406, 06300408 (June 30, 2004; Missoula County, S. Engstrom);
07010401, 07010402, 07010403, 07010404 (July 1, 2004; Missoula County, S. Engstrom);
07070402, 07070403, 07070407 (July 7, 2004; Missoula County, S. Engstrom); 07130401,
444 ... Kuo
07130406, 07130407, 07130408, 07130409 (July 13, 2004; Missoula County, S. Engstrom);
07140401, 07140402, 07140403 (July 14, 2003; Missoula County, S. Engstrom).
IDAHO—06040401 (June 4, 2004; Valley County, K. Greger); 06180423 (June 18,
2004; Valley County, K. Greger); 06190401 (June 19, 2004; Valley County, K. Greger).
OREGON—04070402 (April 7, 2004; Lane County, M. Sheller); 08070404 (August 7,
2004; Jefferson County, M.C. Carter). COLORADO—08310301 (August 31, 2003; La
Plata County, G. Fitzgerald). Accession numbers are from the Morel Data Collection
Project. All collections are housed in the Mycology Collection of the Field Museum of
Natural History (F), Chicago.
COMMENTS — The densely tomentose (and frequently black) surfaces of young
specimens of Morchella tomentosa separate it easily from other species of
Morchella—though older specimens (especially those that have been exposed to
prolonged sunlight) may have eroded ridges and paler colors that approximate
the colors of M. esculenta-like yellow morels, leading to confusion. However,
pale and less tomentose specimens still demonstrate pigmented hairs on the
sterile ridges and stipe surface and can thus be separated microscopically.
Morchella tomentosa is known as the “gray morel” by commercial collectors in
western North America; eastern and Midwestern collectors, however, use the
same term for immature forms of M. esculenta-like yellow morels.
In North America Morchella tomentosa has been described by Pilz et al. (2004)
as “PS D” (putative species D); by Kuo (2005) as the “black foot morel”; by
Kuo (2006) as the “fuzzy foot morel”; and by Pilz et al. (2007) as the “gray
morel.” Additionally, the name M. atrotomentosa has been applied (McKnight
& McKnight 1987) to a western North American burn-site morel that arguably
meets the description of M. tomentosa. However, M. atrotomentosa is an invalid
taxon (McFarlane et al. 2005, Pilz et al. 2007): the basionym was originally
published by Moser (1949) as a temporary (ad interim) taxon. No type collection
was designated for Moser’s “Morchella esculenta (L.) var. atrotomentosa nov. vat.
ad. [sic] int. which Moser (1949) documented fairly rigorously as a burn-site
morel from the Alps that shares many of the morphological features emphasized
for the present species—including the dark, tomentose hymenophore and stipe
and the pigmented hairs. Thus the epithet tomentosa is chosen in part to reflect
potential continuity with Moser’s invalid taxon while also acknowledging that
the North American species, while always tomentose, is not always black-
tomentose. Moser described only a black to blackish brown hymenophore and
stipe.
Jacquetant (1984) described and illustrated “Morchella atrotomentosa (Moser)
Bride.” While the watercolor illustrates a species similar to M. tomentosa, the
description is inadequate for thorough comparison, and Jacquetant does not
document any material examined. Further, no reference is provided for the
Bride authority. Attempts to locate a publication by Bride, or a citation for a
publication by Bride, have been unsuccessful. Moreover, the combination “M.
Morchella tomentosa sp. nov. (North America) ... 445
atrotomentosa (Moser) Bride” is built on an invalid basionym. Validation of
Moser’s name would have required a proposal to conserve —a conspicuous
maneuver that would be easily discovered in standard taxonomic references.
Morchella rufobrunnea in California and Oregon
Morphological analysis and preliminary molecular results (see Kuo 2005, 2006)
indicate that the winter-fruiting yellow morel from West-Coast landscaping
sites is Morchella rufobrunnea Guzman & F. Tapia. This species is quite distinct
and well characterized by its ecology, its abruptly conical young cap with pale
ridges and nearly black pits, and its rufescence.
Guzman & Tapia (1998) described Morchella rufobrunnea based on Mexican
collections from Xalapa, Veracruz, on Mexicos Gulf Coast, made along a
roadbed and in a garden. Collections from landscaping and similar disturbed-
ground sites in California and Oregon matching the M. rufobrunnea isotype
(MDCP 06059601) have been studied (see Specimens Examined, below). All
specimens demonstrate a morphology consistent with the original description
of the species.
Since Morchella rufobrunnea is so visually distinct it can often be recognized in
photographs; the species appears to have been described as “M. deliciosa” and
as “M. esculenta” by western North American authors. Arora (1986) depicts
a “coastal Californian form of Morchella deliciosa” growing “in gardens and
other suburban habitats” that appears to match M. rufobrunnea. Photographs
of California collections of “M. deliciosa” in Wood & Stevens (2007) also match
M. rufobrunnea; the authors describe a mushroom occurring “in bark/wood
chips of fresh landscaping or in disturbed locations, e.g. near compost heaps,
fire pits, dirt basements, logging roads etc.” Additionally, photographs of “M.
esculenta” in Ower (1982) suggest that the species cultivated by Ower (and,
later, others) was M. rufobrunnea. I have examined cultivated morels from
Diversified Natural Products (MDCP 03110601, produced in Michigan) that
also match M. rufobrunnea, and represent an original California collection (G.
Mills, pers. comm.)
SPECIMENS EXAMINED — MEXICO: VERACRUZ—06059601 (June 5, 1996; Xalapa,
G. Guzman; ISOTYPE). UNITED STATES OF AMERICA: CALIFORNIA—03180301
(March 18, 2003; Solano County, W. Andrew); 03300302 (March 30, 2003; Sacramento
County, D. Kimberling); 03180401 (March 18, 2004; Ventura County, R. Sharman);
12110401 (December 11, 2004; San Diego County, C. Nielsen); 03060501 (March 6,
2005; Santa Clara County, M. Munch); 03270501 (March 27, 2005; Siskiyou County, J.
Plischke); 03290501 (March 29, 2005; Tulare County, J. Plischke). OREGON—03080501
(March 8, 2005; Jackson County, J. Petersen). MICHIGAN—03110601 (March 11, 2006;
produced in Mason County, M. Kuo). Accession numbers are from the Morel Data
Collection Project. All collections are housed in the Mycology Collection of the Field
Museum of Natural History (F), Chicago.
446 ... Kuo
Acknowledgments
The author thanks Carol Carter for many years of patient help in understanding morels,
Gastén Guzman for providing isotype material, Kerry O'Donnell, Carol Schmudde, Ron
Petersen, Fred Kranzlin, Harald Kellner, Gary Mills, Sue Yokum, Robert Licking, Kate
Klipp, and Stephen Canfield. Presubmission reviews were kindly provided by Tom Volk,
Rodham Tulloss, Patrick Leacock, and Andrew Methven. The author also thanks the
following individuals for the kind donation of specimens cited in this paper: William
Andrew, Carol Carter, Scott Engstrom, Gretchen Fitzgerald, Ken Greger, John Holmes,
Dan Kimberling, Mary Munch, Carl Nielsen, Jamie Petersen, John Plischke HI, Steve
Schwartzman, Robert Sharman, and Mike Sheller.
Literature cited
Arora D. 1986. Mushrooms demystified: A comprehensive guide to the fleshy fungi. Berkeley,
California, Ten Speed Press.
Guzman G, Tapia F. 1998. The known morels in Mexico, a description of a new blushing species,
Morchella rufobrunnea, and new data on M. guatemalensis. Mycologia 90: 705-714.
Jacquetant E. 1984. Les morilles. Paris, La Bibliotheque des Arts.
Kuo M. 2005. Morels. Ann Arbor, University of Michigan Press.
Kuo M. 2006. MDCP morel taxa. www.mushroomexpert.com/mdcp/results_legend.html
(downloaded on June 27, 2008).
McFarlane EM, Pilz D, Weber NS. 2005. High-elevation gray morels and other Morchella species
harvested as non-timber forest products in Idaho and Montana. Mycologist 19: 62-68.
McKnight KH, McKnight VB. 1987. Mushrooms (Peterson Field Guides). New York, Houghton
Mifflin.
Moser M. 1949. Uber das massenauftreten von Formen der Gattung Morchella auf Waldbrandflachen.
Sydowia, Annales Mycologici ser. II, 3: 174-195.
Ower R. 1982. Notes on the development of the morel ascocarp: Morchella esculenta. Mycologia
74: 142-144.
Pilz D, Weber NS, Carter MC, Parks CG, Molina R. 2004. Productivity and diversity of morel
mushrooms in healthy, burned, and insect-damaged forests of northeastern Oregon. Forest
Ecology and Management 198: 367-386.
Pilz D, McLain R, Alexander S, Villarreal-Ruiz L, Berch S, Wurtz TL, Parks CG, McFarlane E, Baker
B, Molina R, Smith JE. 2007. Ecology and management of morels harvested from the forests of
western North America. Portland, Oregon, USDA General Technical Report PNW-GTR-710.
Wood M, Stevens F. 2007. Morchella deliciosa. http://www.mykoweb.com/CAF/species/Morchella_
deliciosa.html (downloaded on June 27, 2008).
MYCOTAXON
Volume 105, pp. 447-454 July-September 2008
Ganoderma vivianimercedianum sp. nov.
and the related species, G. perzonatum
MABEL GISELA TORRES- TORRES? 2*, LAURA GUZMAN-DAVALOS?
* magitoto@yahoo.com
' Universidad Tecnologica del Choc6, Ciudadela Medrano
Quibdo, Choco, Colombia
lguzman@cucba.udg.mx
? Departamento de Botanica y Zoologia, Universidad de Guadalajara
Apartado postal 1-139, Zapopan, Jal., 45101, Mexico
&
ADRIANA DE MELLO GUGLIOTTA?
agugliottaibot@yahoo.com.br
° Instituto de Botanica,Secdo de Micologia e Liquenologia
Caixa Postal 3005, CEP 01061-970, Sao Paulo, SP, Brazil
Abstract — A new species of Ganoderma, G. vivianimercedianum is described from
Cuba, Brazil and Mexico. It is distinguished by its flabelliform basidioma with a
contracted base and a conchate, radially rugose pileus with claviform and granulated
cuticle cells and context that is not completely homogeneous. Comparisons with related
species and a full description of G. perzonatum are also provided.
Key words — Ganoderma argillaceum, G. resinaceum
Introduction
In the course of the study of Ganoderma developed by the first author,
two specimens deposited in NY, both marked as representing the type of
G. argillaceum Murrill, were examined. However, these two specimens differ
in several features, raising questions both about their conspecificity and the
species concept of G. argillaceum. Moncalvo & Ryvarden (1997) cite the usually
accepted type of G. argillaceum, NY collection ES. Earle 658 (5 July 1904),
which corresponds to the characteristics described in the protologue and is
the only specimen mentioned by Murrill (1908). The other specimen appears
more closely related to G. perzonatum and G. resinaceum Boud. The types or
authentic material of G. perzonatum and G. resinaceum were compared with
the material in question.
448 ... Torres-Torres, Guzman-Davalos & Gugliotta
As a result of these type studies, we propose and describe a new species based
on the second collection previously referred to G. argillaceum and additional
specimens from Brazil and Mexico. We also provide an updated description of
G. perzonatum.
Materials and methods
Materials studied are deposited at BPI, ENCB, IBUG, NY and SP. The colors
are based in Kornerup & Wanscher (1963). Micromorphological observations
were made from basidiome tissues mounted in 5% KOH and Melzer’s reagent.
Basidiospore shape is based on the Q coefficient (length-width, Bas 1969) of 20
randomly selected spores.
Taxonomy
Ganoderma vivianimercedianum Torres-Torres, sp. nov. Figs. 1-10
MycoBank MB511309
Fructificatio lignicola, perenne, sessilia vel substipitata; superficies demum rubro-brunnea,
crustosa vel laccata; pori 3-5 per mm; contextus pallido-bubalinus vel purpureo-brunneus,
non zonae resinosa; systema hypharum trimiticum; hyphae generatoriae fibulatae;
hyphae skeletales nonseptatae, arboriformes; cutis hymeniodermiformis, pallide brunneae,
dextrinoidea, elementis entire vel lobata, 36-72 x 7.2-13.6 um; basidiosporae ellipsoideae,
truncate, luteo-brunnea, paries duplexis, cum columnae interparietis, 9.6-11.2 x 6.4-8
um.
MEXICO, Estado de México, Valle del Tepeite, 10 km NE of Santa Maria, 10 August
1986, E. Bastidas- Varela s.n (Holotype: ENCB).
EtyMo_Loey: In memory of Angie Viviana and Rosa Mercedes, niece and aunt of the
first author.
BASIDIOMATA 6.5-10 x 8.5-13 x 1.3-1.5 cm, annual, sessile to substipitate,
then with a contracted base, single, woody-corky consistency, light in weight.
Piieus flabelliform, conchate; surface glabrous, bumpy, concentrically sulcate,
radially rugose, glossy; with a laccate crust, not cracking, difficult to remove,
easy to penetrate with fingernail; violet-brown (11F7-11F8) in the 80 to 90% of
the surface, to henna (7E8-8E8) in the periphery; margin whitish to yellowish,
generally entire to slightly lobulate, thin, smooth. SussTipE 2 x 1.3 cm,
lateral, cylindrical, solid, surface shiny, darker than pileus, red-wine to almost
black. CONTEXT 0.9-1.3 cm thick, 0.6 cm average, fibrous-corky, zonate, not
completely homogeneous, caramel (6C6) above to dark brown (7F7) toward
the tubes; with resinous incrustations (not observed in the type). PoREs 3-5 per |
mm, angular to circular, woody; pore surface pastel-yellow (2A4), darkening to
brown (6D8) when bruising or aging; tubes 0.5-0.9 cm thick, up to 1.2 cm in
the base, unstratified, concolorous with the inferior part of the context.
HYPHAL SYSTEM trimitic. CONTEXTUAL TRAMA with generative hyphae 1.8-
4.8 um diam, hyaline, branched; skeletal hyphae 3.2-12.4 um diam., generally
Ganoderma vivianimercedianum sp. nov. ... 449
Figs. 1-3. Ganoderma vivianimercedianum basidiomata.
1: holotype., 2: Pacheco s.n., 3: Sotao et al. 88.21.26. Scale bar = 1 cm.
solid to thick-walled, non-septate, arboriform, yellowish to golden-yellow,
predominant; binding hyphae 1.9-8.8 um diam., solid, non-septate, yellowish
to light yellowish-brown. HyYMENOPHORAL TRAMA as contextual trama.
PILEIPELLIS with cuticle cells 36-65 x 7.2-12 um, narrowly claviform to clavate,
450 ... Torres-Torres, Guzman-Davalos & Gugliotta
Figs. 4-6. Ganoderma vivianimercedianum basidiomata showing context that is not completely
homogeneous. Arrows indicate resinous incrustations. 4: Earle s.n., 5: Pacheco s.n., 6: Sotao et al.
88.21.26. Scale bar = 1 cm.
entire or with up to two lateral protuberances, not branched or few with two
apical branches; thick-walled, unstratified, golden-yellow, with granulations
in the apex; cells dextrinoid in Melzer’s reagent. Bastp1osporEs 8.8-11.2(-12) °
x 6.4-8 um, Q = 1.25-1.5, ellipsoid, apex truncate, light yellowish-brown,
negative in Melzer’s reagent; perisporium wrinkled, hyaline; exosporium with
inter-walled pillars up to 0.3 um thick, subfree; endosporium wrinkled, with
apical germ pore. Basidiospores print rust-brown (6E8) (from the deposit of
spores on the pileus). BAsIDIA not seen. CysTrp1A absent.
Ganoderma vivianimercedianum sp. nov...
Figs. 7-10. Microscopy of Ganoderma vivianimercedianum (Earle s.n).
7: Cuticle cells, 8: basidiospores, 9: binding hyphae, 10: skeletal hyphae.
Scale bar = 8 um.
SUBSTRATA — Dead wood.
Hasitat — Secondary tropical forest, mangrove.
SPECIMENS STUDIED - CUBA, sine data (NY). BRAZIL, Guaiba, Rio Grande do Sul,
Fazenda da Faculdade de Agronomia e Veterinaria, em lenho morto, 29 March 1963, J.P.
de Costa-Neto s.n. (SP70533); Sao Paulo, Tremembé da Cantareira, Villa Amalia, Horto
Forestal SPSF, podridao da madeira, 20 May 1952, C.D.E. Pacheco s.n. (SP109563);
Amapa, Ilha de Maraca, tronco de madeira en descomposi¢ao, 24 October 1988, H.
Sotao et al. 88.21.26 (SP233297). MEXICO, Estado de México, Valle del Tepeite, 10 km
NE of Santa Maria, 10 August 1986, E. Bastidas-Varela s.n. (holotype, ENCB); Jalisco,
Zapopan, Centro Universitario de Ciencias Bioldgicas y Agropecuarias, Departamento
de Madera Celulosa y Papel, on dead wood, 21 October 2006, M.G. Torres-Torres 699
(IBUG); Veracruz, 5 km from Jaltipan, on dead wood, 19 March 1978, J. Pérez-Ortiz
1278 (ENCB).
.451
452 ... Torres-Torres, Guzman-Davalos & Gugliotta
Ganoderma perzonatum Murrill, North Amer. Flora 9: 121, 1908. Figs. 11-13
BASIDIOMATA 2:5-8.5 x 4.8 x 0.7-1.5 cm, annual or perennial, substipitate, single
to imbricate, woody-corky consistency, light in weight. Piteus flabelliform;
surface glabrous, bumpy, glossy, concentrically sulcate; with a laccate crust,
not cracking, difficult to remove, easy to penetrate with fingernail; violet-
brown (11F8) almost homogeneous grading to garnet-red (11E8) towards the
periphery; margin concolorous with pileus, entire, thin, smooth. CONTEXT
0.3-0.9 cm thick, 0.7 cm average, fibrous, zonate, not completely homogeneous,
pale-orange to light-orange (5A3, 5A4) above, to light brown (6D7) toward
the tubes; with resinous bands. Porgs 4-5 per mm, angular to circular, woody;
pore surface pale-yellow (2A3); tubes 0.5-0.9 cm thick, up to 1.2 cm in the
base, unstratified to stratified, concolorous with the inferior part of the context.
HYPHAL SYSTEM dimitic. CONTEXTUAL TRAMA generative hyphae not observed;
skeletal hyphae 6.4-13.6 um diam., generally solid to thick-walled, non-
septate, arboriform, yellowish. HYMENOPHORAL TRAMA as contextual trama.
PILEIPELLIS with cuticle cells 70.4-97.6 x 5.6-20.8 um, cylindrical, apex obtuse
to rarely subcapitate, entire or with one lateral protuberance; thick-walled,
generally with many layers in the apex, golden-yellow, with concentric elongate
granulations in the apex; dextrinoid with Melzer’s reagent. BASIDIOSPORES 8.4—
10.4 x (6-) 6.4-7.2 (-7.6) um, Q = (1.25-) 1.36 -1.53, ellipsoid, apex truncate, light
yellowish-brown, negative in Melzer’s reagent; perisporium wrinkled, hyaline;
exosporium with inter-walled pillars up to 0.3 um thick, free; endosporium
wrinkled, with apical germ pore. BAsipIA not seen. CysTIDIA absent.
cece
ane
Soh
Steere
rn ——— - aos a
Se NST oO 2 CO IPR RE IIAeARSPLITIE
Figs. 11-13. Microscopy of Ganoderma perzonatum (Lectotype).
11: basidiospores, 12: cuticle cells, 13: skeletal hyphae. Scale bar = 8 um.
Ganoderma vivianimercedianum sp. nov. ... 453
CoMMENTS — The lectotype is a half basidioma in good condition. Fragments
of the type in BPI consist of half of one small imbricate basidioma, two ~6 x
2.5 cm fragments, and two slices — one including pileus, context and tubes and
the other with the laccate crust in both surfaces (without hymenophore) and
context.
SUBSTRATA — Mango log.
Hasitat — Not specified.
SPECIMEN STUDIED — CUBA, Santiago de Las Vegas, 8 November 1904, ES. Earle 309
(Holotype: NY; Isotype: BPI).
Discussion
The type of G. argillaceum deposited in NY is a specimen marked as: CUBA,
Province of La Habana, Santiago de Las Vegas, on dead mango log, 5 July 1904,
ES, Earle 658.
The second specimen in NY also annotated as type of G. argillaceum [“part of
type, Earle, Cuba’ (hand written label)] was collected by the same collector
and presumably from the same locality; no number or other data is available.
Moncalvo & Ryvarden (1997) did not refer to this collection, but J.E Wright,
who did examine the specimen, wrote (1967, hand written label): “Very much
like G. perzonatum Murr.!” M.E. Bazzalo also examined the specimen, which
he referred to G. resinaceum (1980, hand written label). After examining the
specimen ourselves, we agree with Wright that the material does not correspond
to G. argillaceum but is morphologically related to G. perzonatum. However, it
also differs from G. perzonatum in several features; consequently, we describe
it above as a new species.
Ganoderma vivianimercedianum can be recognized by the flabelliform
basidioma with a contracted base, light brown and not completely homogenous
context, and the narrowly claviform to clavate cuticle cells with scattered
granulations concentrated at the apex. The Brazilian and some Mexican
specimens have resinous incrustations in the context near the pileus base that
were not observed in Earle’s specimen deposited in NY (which does, however,
lack the very base).
TABLE | compares the main differences among G. vivianimercedianum and the
morphologically similar G. argillaceum, G. perzonatum, and G. resinaceum.
In general, Ganoderma argillaceum has larger (10.4-13.6 x 7.2-9.6 tm)
basidiospores and cuticle cells without granulations. Ganoderma perzonatum
has slightly smaller (8.4-10.4 x 6.4-7.2 um) basidiospores, remarkable large
cylindrical cuticle cells with concentric elongate granulations present only
at the apex, as mentioned above. Ganoderma resinaceum has basidiospores
454 ... Torres-Torres, Guzman-Davalos & Gugliotta
similar in size (11.2-13.6 x 6.5-7.4(-8.1) um) to G. argillaceum (and larger than
G. perzonatum), narrowly clavate, almost cylindrical cuticle cells, and a context
that is fibrous-spongy, homogeneous, light reddish-brown, and lacking resinous
bands. Ryvarden (2004) suggested that G. perzonatum and G. resinaceum might
be related, but we consider these as two independent species, each one with
unique features.
Table 1. Morphologic features of Ganoderma argillaceum, G. perzonatum,
G, resinaceum and G. vivianimercedianum.
SPECIES CONTEXT BASIDIOSPORES nities GRANULATIONS
CELLS
G. argillaceum not completely —_19.4-13.6 x clavate no
homogeneous 7.2-9.6 um
G. perzonatum not completely 8.4-10.4 x cylindrical yes
homogeneous (6-)6.4-7.2(-7.6) um
G. resinaceum homogeneous 11.2-13.6 x clavatealmost —_yes
6.5-7.4 (-8.1) um cylindrical
G, vivianimercedianum not completely 8.8-11.2 (-12) x narrowly yes
homogeneous 6.4-8 um claviform to
clavate
Acknowledgments
We extend our thanks to Tatiana Baptista Gibertoni, Cony Decock and Gast6n Guzman
who kindly reviewed the manuscript. We are grateful to ENCB and NY curators for
the loan of materials. Also we acknowledge to Servando Carvajal for reviewing the
Latin diagnosis. The first author thanks to the Red Latinoamericana de Botanica for
the improvement scholarship (RLB-05-P5) for her stay in the Instituto de Botanica
da Secretaria de Estado do Meio Ambiente, Sao Paulo. She also thanks Universidad
Tecnoldgica del Chocé and Project NUFFIC-Alterra, Wageningen University for grants
for her stay in Mexico. Adriano Spielmann is thanked to take the pictures of Brazilian
specimens (SP). Funds were also obtained from CONACYT (project CONACYT-SEP-
2003-C02-42957) and Universidad de Guadalajara (project 62935).
Literature cited
Bas C. 1969. Morphology and subdivision of Amanita and a monograph of its section Lepidella.
Persoonia 5: 285-579.
Kornerup A, Wanscher JH. 1963. Methuen handbook of colour. Methuen, London. ‘
Moncalvo JM, Ryvarden L. 1997. A nomenclatural study of Ganodermataceae Donk. Synopsis
Fungorum 11, Fungiflora, Oslo, Norway. 114 p.
Murrill WA. 1908. Polyporaceae. North American Flora 9: 1-131.
Ryvarden L. 2004. Neotropical Polypores, part 1, Synopsis Fungorum 19, Fungiflora, Oslo, Norway.
229p.
MYCOTAXON
Volume 105, pp. 455-468 July-September 2008
New records and range extensions in the
North American lignicolous lichen flora
TOBY SPRIBILLE
tspribi@uni-goettingen.de
Albrecht von Haller Institute of Plant Sciences, University of Gottingen
Untere Karspiile 2, D-37073 Gottingen, Germany
CurTIs R. BJORK
Stillinger Herbarium, University of Idaho
Moscow, ID, 83843, U.S.A.
Abstract — Based on recent intensive studies of wood-dwelling lichens especially in
western North America, we report a series of new records and range extensions. Six
species are reported as new to North America: Arthonia ligniariella, Elixia flexella,
Gyalideopsis helvetica, Lecidea scabridula, Lecidea symmictella and Xyloschistes
platytropa. Chaenothecopsis nigra is again confirmed for North America. Buellia arborea,
B. chloroleuca, Catillaria erysiboides, and Lignoscripta atroalba are new to Canada and
Lecidea pullata and Xylographa trunciseda are new to eastern North America. We also
provide range extensions for all four species of Pycnora known from North America.
In all we include new records from Alaska, Arizona, British Columbia, Colorado,
Idaho, Manitoba, Montana, New Brunswick, Newfoundland, North Dakota, Northwest
Territories, Nunavut, Ontario, Québec, Washington and the Yukon.
Key words — calicioid, crustose lichens, floristics, Pacific Northwest, saproxylic, wood
Introduction
The study of crustose lichens in North America is enjoying something of a
renaissance. In the last 10 years, hundreds of species of crustose lichens have
been reported as new to science or new to North America as a result of projects
such as the Sonoran Lichen Flora (Nash et al. 2002, 2004, 2007) and intensified
inventory efforts in eastern North America (e.g., Harris & Lendemer 2005)
and the Pacific Northwest/western Canada (e.g., Printzen & Tonsberg 1999,
Spribille et al. 2008a). The fact that the rate of new descriptions and first records
is not letting up belies the potential depth of the well of lichen diversity in
North America.
456 ... Spribille & Bjork
As part of a broad-scale study of lignicolous lichens (Spribille et al. 2008b), we
have accumulated a large number of records of previously unknown or poorly
documented species, mainly from northwestern North America but also
from other regions. While not every record reported here necessarily comes
from wood, the general purpose of the present paper is to document a series
of significant new records and range extensions of predominantly lignicolous
lichens.
Materials and methods
Apothecia and thallus were investigated by light microscopy on hand-cut
sections, mounted in water and/or 10% KOH (hereafter K), and stained with
Nees (Merck 1.09261; hereafter IKI) or lactophenol cotton blue (Merck 1.13741).
A polarizing (POL) filter was used to detect birefringent crystals. Thin layer
chromatography (TLC) was carried out according to the methods described by
Culberson (1972) and Culberson & Johnson (1982).
Results and discussion
Arthonia ligniariella Coppins
Arthonia ligniariella was described from Scotland by Coppins (1989) and
has been reported since apparently only from Sweden (Gustavsson 1995, Thor
2008). It was originally thought to be an obligate lignicole. In British Columbia,
the species was found on bark of Thuja plicata, a habitat that often supports
species that are otherwise lignicoles. Its occurrence here was briefly mentioned
by Houde et al. (2007), who also provided a description of the broader habitat
setting. A. ligniariella can be easily recognized on account of its non-amyloid
hymenium, a trait that is rare in Arthonia and smaller ascospores than e.g.
A. ligniaria (Coppins 1989). It is new to North America.
SPECIMEN EXAMINED: CANADA. British Columbia. Rocky Mountains, near border with
Alberta, Albert River, 50°37.693’N, 115°32.973’W, in old-growth Thuja plicata forest, on
underhang of Thuja, 1224 m, 30 Jul 2005, Spribille 16780 & Houde (hb. Spribille, ver.
B. Coppins).
Arthonia spadicea Leight.
This species was originally described from Britain and has been reported from
Germany and elsewhere in Europe as well as Japan (Kashiwadani & Thor 1995).
In North America the species has been reported from the Great Lakes region -
(Hedrick & Lowe 1936, Thomson 1951, Wetmore 1981) and North Carolina
(Schmitt & Slack 1990). In British Columbia the species was found on wood of
a snag in old-growth Tsuga heterophylla forest. It may be genuinely rare here,
as extensive collecting of this habitat in the inland rainforests of east-central
British Columbia has so far yielded only this one specimen. Arthonia spadicea
Se ————— a ee ee eee
New lichen records in North America ... 457
is characterized by its 1-septate ascospores 9.5-11 x 4-4.5 um with unequal
cells, presence of K+ violet hymenial pigments, and presence of Trentepohlia
as photobiont. The British Columbia specimen compares well with European
material; it is new to western North America.
SPECIMEN EXAMINED: CANADA. British Columbia. Selkirk Mountains, Incomappleux
River drainage, Boyd Creek, 50°51.830°N, 117°31.636’W, 1185 m, lignicolous on soft
snag in old-growth Tsuga heterophylla forest, 23 Aug 2005, Spribille 17810 & Pettitt
(CANL).
Buellia arborea Coppins & Tonsberg
Buellia arborea was first reported for North America by Tonsberg & McCune
(2001) and until now has been known only from this one published record, from
western Montana. The species is, in fact, not uncommon on wood in dry, low
elevation Pinus ponderosa forests of the inland Pacific Northwest and British
Columbia but may evade detection because it can only be reliably identified
using TLC. It can easily be confused with Buellia griseovirens and Xylographa
vitiligo s.lat., but differs from both species in having more consistently concave
soralia flecked in a ‘salt-and-pepper’ fashion with darker soredia and in the
lack of stictic or norstictic acid (instead: atranorin and pseudoplacodiolic
acid; Tonsberg 1992). We report it here as new to Canada and, in a significant
southward range extension, from Arizona.
SPECIMENS EXAMINED: CANADA. British Columbia. East Kootenays, NE of Kimberley,
Ta Ta Creek, 49°50.823’N, 115°47.723’W, 792 m, on decorticated log in open xeric Pinus
ponderosa woodland, 4 Aug 2005, Spribille 17166 (hb. Spribille), 17167 (CANL, hb.
Spribille); U.S.A. Arizona. Graham Co., Pinaleho Mountains, Shannon Campground,
32°41°N, 109°51’W, on decorticated tree, Pseudotsuga forest, 24 Sep 1977, Nash 14461
(ASU).
Buellia chloroleuca Korb.
Buellia chloroleuca is one of the most common fertile species of Buellia on
logs in the northern Rocky Mountains. It was first reported for North America
by Bungartz et al. (2007) after long having passed in North America literature
under the name Buellia zahlbruckneri (for a discussion of the nomenclatural
problems, see Bungartz et al. 2007). It is otherwise known from Europe (e.g.,
Nordin 2000). It can be recognized by its large Callispora-type ascospores which
are 16-22 x 7-9.5 um, and the presence of the xanthone 6-O-methylarthothelin,
usually in concentrations sufficient to give a C+ orange reaction. Nordin
(2004) placed it in the genus Tetramelas, but that genus has not been widely
recognized. The predominantly lignicolous habit of B. chloroleuca in western
North America and southern Europe contrasts with its corticolous habit in
northern Europe (Nordin 2000), an ecological difference that deserves further
investigation. We report it here as new to Canada (British Columbia, Yukon),
Idaho, Montana and Washington.
458 ... Spribille & Bjork
REPRESENTATIVE SPECIMENS EXAMINED (>20 seen): CANADA. British Columbia.
Rocky Mountains, NE of Canal Flats, Dry Creek, 50°15°19"N, 115°42°53”W, 1000-1100
m, on rotten log in 30-40 yr old clearcut, 26 Jul 2004, Spribille 14779 (hb. Spribille, det.
A. Nordin, 2004, also !Bungartz); ibid., E of Canal Flats, Moscow Road, 50°14.908’N,
115°30.020’W, 1182 m, 26 Jul 2005, Spribille 16507 (UPS); ibid., Spribille 14779 (CANL);
East Kootenays, NE of Kimberley, Ta Ta Creek, 49°50.823’N, 115°47.723'W, 792 m, on
decorticated log in open xeric Pinus ponderosa woodland, 4 Aug 2005, Spribille 17165
(CANL); Yukon. Carpenter Lake, Wernecke Mountains, 64°30’N, 135°06’W, dwarf birch
and willow communities, with scattered clumps of white spruce, 25 Jul 1972, Scotter
19376 (CANL, filed under Xylographa abietina); along Alaska Highway S of Johnson's
Landing, near Teslin Lake, 60°28.641’N, 133°16.527’°W, 719 m, on rotten log, 8 Oct
2007, Spribille 25076 (ALA); U.S.A. Idaho. Bonner Co., Selkirk Mountains, trail to
Upper Priest Lake, 48°45’N, 116°51’W, on wood of conifer log, Jul 2006, Bjork 13048
(UBC); Montana. Flathead Co., near confluence of Lime and Magnesia Creeks, Collins
Ranch, 48°38.793’N, 114°53.180’W, 1011 m, on rotten log, edge of Picea swamp, 20 Jul
2006, Spribille 20313 & Pérez-Ortega (ALA, BG); Lincoln Co., W of Eureka, Black Butte,
48°52.151’N, 115°06.701’W, 974 m, on log, 31 Jul 2006, Spribille 21240 & Pérez-Ortega
(hb. Spribille); ibid., Salish Mountains, Weigel Creek, 48°28.676'N, 114°58.905’W, 1147
m, on wood, 10 Aug 2006, Spribille 21944 (BG); Washington. Douglas Co., Badger
Mountain, 47°36’N, 120°08’W, on log of Pinus ponderosa, Aug 2005, Bjork 12188
(UBC).
Catillaria erysiboides (Nyl.) Th. Fr.
This species was first reported for North America by Printzen & Tonsberg
(1999), based on the 19" century Wilhelm Suksdorf collection described by
Willey (1887) as Arthonia carneorufa. There are no other published records
for North America to our knowledge. We have found the species sporadically,
primarily on the sides of rotten logs in dry forests, in areas where grass or herbs
grow up around the logs, rarely on the tops of logs, other specimens have been
gathered from vertical wood surfaces on snags and stumps, always near ground
level. The species can easily be recognized by its combination of 1-septate
ascospores with one cell larger than the other (likely why Willey assigned it to
Arthonia), the presence of a well developed excipulum, and the ‘Porpidia-type
ascus. We report it here as new to Canada.
SPECIMENS EXAMINED: CANADA. British Columbia. Rocky Mountains, E of Canal
Flats, Moscow Road, 50°20.825’N, 115°35.886'W, 1069 m, 1 Aug 2005, Spribille 16938
(hb. Spribille); East Kootenays, E of Canal Flats, Kootenay River valley, Dry Creek,
50°15.120°N, 115°42.693’W, 1122 m, on decorticated log, 4 Aug 2005, Spribille 17206
(CANL); Thompson Plateau, Opax Mountain silvicultural study site, 20 km NW of
Kamloops, 50°49°N, 120°28’W, on wood of snag near ground-level, Sep 2007, Bjork
15094 (UBC).
Chaenothecopsis nigra Tibell
This species occurs on wood, over thalli of Chaenotheca species or on groups
of free-living algae and has been reported from New Zealand (type locality),
South America, Great Britain, Lithuania, Poland, Russia, Sweden and Japan
New lichen records in North America ... 459
(Titov 2006). Rikkinen (2003) reported Chaenothecopsis nigra from Oregon
with ‘cf’ on account of deviations in ascospore size. It was also reported, with
a question mark, by Harris & Lendemer (2005), although the two characters
cited by the latter (dark ascospore septum, but dark greenish KOH- stalk)
could also lead to other Chaenothecopsis species. Titov (2006) provided the first
confirmed record from North America, from Mt. Cain on Vancouver Island.
We report the first confirmed specimen here for the United States.
SPECIMEN EXAMINED: U.S.A. Idaho. North Fork Clearwater River drainage, Isabella
Creek, 46°52’N, 115°37’W, on cubed cracked wood on leaning snag, 8 Jul 2001, Spribille
11543-B & Hauck (UPS, det. L. Tibell).
Cladonia botrytes (K.G. Hagen) Willd.
Cladonia botrytes is one of the few obligately lignicolous macrolichens
(Spribille et al. 2008b). Although the species is well known in the boreal zone
of North America and has been reported from British Columbia and Alaska, it
has been reported south of Canada in the west only from Montana (DeBolt &
McCune 1993), where it is locally common in Flathead and Lincoln Counties.
We report the species here as new to Washington.
SPECIMEN EXAMINED: U.S.A. Washington. Pend Oreille Co., Frater Lake, 48°39.266'N,
117°29.090'W, 968 m, on stump, 19 Aug 2005, Spribille 17479 (WTU).
Elixia flexella (Ach.) Lumbsch
Elixia flexella was long treated within the genus Lithographa but was
recognized as a distinct, monospecific genus by Lumbsch (1997). The species
was described from France and subsequently reported from the United
Kingdom (Hawksworth et al. 1980), Austria and Italy (Lumbsch 1997), eastern
Europe (e.g., Miadlikowska 1993), Scandinavia (Santesson et al. 2004), the
Komi Republic, western Russia (Hermansson et al. 2006) and Japan (Kurokawa
& Kashiwadani 2006). It can be recognized by the distinctive folded, involute
exciple that often gives the mature apothecia a triangular or square shape, IKI+
reddish hymenium, and simple ascospores 6-8 x 3-4.5 um. It is new for North
America.
SPECIMENS EXAMINED: CANADA. British Columbia. Southern interior, Sicamous Creek
Silvicultural Research Site, 50°49’N, 118°50’W, on snag, north-facing slope, subalpine
forest of Abies lasiocarpa and Picea engelmannii, 27 Jul 1997, Goward 97-393 (UBC, det.
G. Thor); Cariboo Mountains, Wells Gray Provincial Park, shortly S of the park road
immediately north of Second Canyon Creek, 51°46.2’N, 120°0.6’W, weathered wood of
a Pseudotsuga stump, 27 Feb 2006, Bjérk 12191 (UBC); U.S.A. Idaho. Shoshone Co., St.
Maries River Valley, 47°3.5°N, 116°17.9'W, swamp forest shortly east of Hwy. 3 between
Clarkia and Emerald Creek, on bark of Thuja trunk, 11 Apr 2008, Bjork 15312 (UBC);
Washington. Skamania Co., Columbia River Gorge, 45°42.9’N, 120°49.2’W, on wood of
a Pseudotsuga snag, 19 Mar 2007, Bjork 14088 (UBC).
460 ... Spribille & Bjérk
Gyalideopsis helvetica van den Boom & Vézda
This distinctive species was first described from Austria by van den Boom
& Vézda (2000), and remains one of the rarely reported lignicolous lichens,
being known only from Switzerland, Germany and Austria (Hafellner et al.
2005) and Sweden (Hermansson et al. 2008). When sterile it is difficult to
recognize as the thallus consists only of a thin film interrupted by patches of
concave goniocystangia (van den Boom & Vézda 2000). Fertile specimens are
more easily recognized by the translucent, ‘gummy bear-like’ apothecia and
distinctive muriform ascospores. The species has only been found once during
our field studies of lignicolous lichens, and, fortunately, it was fertile. It is new
to North America.
SPECIMEN EXAMINED: CANADA. British Columbia. East Kootenays, E of Canal Flats,
Kootenay River valley, Dry Creek, 50°15.125’N, 115°42.725’W, 1126 m, on decorticated
log, 4 Aug 2005, Spribille 17193 (hb. Spribille, mixed with Absconditella lignicola).
Hypocenomyce anthracophila (Ny1.) P. James & Gotth. Schneid.
This species is widely reported for the Great Lakes region, the American
Southwest (e.g., Timdal 2002) and South Dakota (Wetmore 1967) and is also
mapped for Alberta, Manitoba and California by Brodo et al. (2001). It was first
reported for northwest North America by Goward et al. (1994) from British
Columbia, but it appears to be rare in the northwest of the continent. We report
it here as new to Washington.
SPECIMEN EXAMINED: U.S.A. Washington. Skamania Co., Columbia River Gorge,
45°42.9°N, 120°49.2’W, on wood of a Pseudotsuga snag, 100 m, 19 Mar 2007, Bjork
14086 (UBC).
Lecidea pullata (Norman) Th. Fr.
Lecidea pullata was first reported for North America by Goward & Thor
(1992) based on a specimen collected in coastal British Columbia. It occurs on
a wide array of bark and wood substrates and is much more common than the
few records would seem to indicate. We report Lecidea pullata as new to the
United States (Colorado, Montana) and eastern Canada (Québec).
REPRESENTATIVE SPECIMENS EXAMINED (>20 seen): CANADA. Québec. Gaspé West
Co., Mont Albert, La Potatiere T.P., on decaying wood, 1971, Muhle s.n. (CANL, filed
under Xylographa abietina); U.S.A. Colorado. Clear Creek Co., N face of Squaw Mountain
along Squaw Pass Road, 39°41’30"N, 105°30’W, on wood, 16 Aug 1983, Nash 22134
(ASU, filed under Xylographa parallela and mixed with Pycnora sorophora); Montana.
Lincoln Co., northern Salish Mountains, Cliff Creek, 48°35’30"N, 115°06’W, on Populus
balsamifera bark, 17 Sep 1996, Spribille 6376 (BG, hb. Spribille; det. T. Tonsberg).
Lecidea scabridula Hedl., nom. illegit., non Mill. Arg.
Lecidea scabridula is known only from a set of type material collected by
Johan Theodor Hedlund in Sweden (Hedlund 1892). Printzen (1995) reported
New lichen records in North America ... 461
that the type material had been studied by Brian Coppins and tentatively
assigned to the order Agyriales based on ascus characteristics. The species
possesses a highly unique set of characteristics: the ascus is quite similar to
the ‘Agyriales-type’ (but see Lumbsch et al. 2007), ascospores are simple, 7-12
x 4-5 um and the exciple and epihymenium are carbonized. However, unlike
in Ptychographa, another genus with ‘Agyrialean’ ascus type and carbonized
exciple, the apothecia are rounded, not lirellate. The systematic position of the
species is the subject of an ongoing study (Spribille & Lumbsch, in prep.), in
which a new name will be created to replace the currently illegitimate one. In
the interim, we report it here as new to North America as a voucher for the
inclusion of the species by Spribille et al. (2008b).
SPECIMENS EXAMINED: CANADA. British Columbia. East Kootenays, White River
drainage, SE of Canal Flats, 50°14.522’N, 115°29.283’W, 1164 m, on decorticated log
with Placynthiella icmalea and Trapeliopsis flexuosa, 28 Jul 2005, Spribille 16631 (hb.
Spribille); ibid., Kootenay River drainage NE of Canal Flats, 50°15.159°N, 115°42.517°'W,
1069 m, on decorticated log with Xylographa vitiligo, Ochrolechia gowardii, Lecidea
rubrocastanea, 5 Aug 2005, Spribille 17249 (hb. Spribille); Northwest Territories. Vicinity
of Glacier (Brintnell) Lake, near camp, 62°5’N, 127°35’W, dead wood on ground, 18 Aug
1939, Raup 3557 (CANL, FH, both filed under Xylographa parallela); U.S.A. Washington.
Okanagan Co., Okanogan National Forest, trail to Roger Lake, off Forest Service Rd. 39,
48°39°30”N, 119°57°50”W, ca. 1775 m, on conifer wood, 14 Oct 1993, Ryan 30733 (ASU,
filed under Xylographa abietina).
Lecidea symmictella Ny].
Lecidea symmictella is a widespread though seldom collected lignicolous
species in northern (Foucard 2001) and central Europe (van den Boom 2002).
Contrary to the suggestion of the name, the species bears little resemblance
to Lecanora symmicta. L. symmictella is characterized by its small apothecia
0.1-0.3 mm diam., simple narrowly ellipsoid ascospores 3-8 x 1.5-2.65 um,
+Micarea-type ascus (not Catillaria-type as stated by Foucard 2001), and the
presence of a POL+ granular epihymenium (Z. Palice, pers. comm.). In North
America, the species may belong to a continental boreal element. It is new to
North America (Montana and Ontario).
SPECIMENS EXAMINED: CANADA. Ontario. Thunder Bay District, Pine Portage, 23
miles N of Nipigon, 49°20’N, 88°20’W, boreal Picea mariana-Abies forest, on rotting log,
26 Aug 1965, Brodo 6137 (CANL, filed under Xylographa abietina); U.S.A. Montana.
Flathead Co., Salish Mtns., near confluence of Lime and Magnesia Creeks, Collins
Ranch, 48°38.793’N, 114°53.180’W, 1011 m, on hard decorticated log, 20 Jul 2006,
Spribille 20251-A & Pérez-Ortega (hb. Spribille).
Lignoscripta atroalba B.D. Ryan & T.H. Nash
This lichen was described as a new genus and species by Ryan (2005),
differing from Xylographa on account of its intense greenish, HNO,+ reddish
epihymenial pigments, pruinose exciple, and bacilliform conidia. At the time,
462 ... Spribille & Bjérk
only a single specimen from Arizona was reported. We report it here as new to
Canada (British Columbia) and the U.S. states of Montana and North Dakota.
SPECIMENS EXAMINED: CANADA. British Columbia. East Kootenays, W of Invermere,
Toby Creek canyon, on wood of abandoned bridge, 20 Jul 2004, Spribille 15316 &
Goward (hb. Spribille). U.S.A. Montana. Lincoln Co., 1 km N of Rexford, 48°54.683’N,
115°09.629'W, 800 m, on wood of old guard rail, 30 Jul 2006, Spribille 21011 (hb.
Spribille); Teton Co., NW of Choteau, on decorticated Pinus flexilis branches, 26 Jul 2006,
Spribille 20668 & Pérez-Ortega (F); North Dakota. Billings Co., Theodore Roosevelt
National Park, South Unit, at Badlands Overlook (3.5 mi E of Medora); Sec. 20, T140N,
R101W, on north facing slope with junipers, on juniper log, 25 Jul 1982, Wetmore 45080
(ASU); McKenzie Co., Theodore Roosevelt National Park, N Unit, on south side of
Cedar Canyon (14 mi SSW of Watford City), Sec. 25, T148N, R100W [added in pencil:
47°36'45°N, 103°22’54”W]; on north facing slope with juniper and few ash, on juniper
log, 20 Jul 1982, Wetmore 44812 (CANL).
Micarea melaena (Nyl.) Hedl.
This is one of the most widely reported epiphytic species of Micarea, being
known from Eurasia (Czarnota 2007), Japan (Kurokawa & Kashiwadani 2006)
and eastern North America (e.g., Wetmore 1981, Harris & Lendemer 2006)
and in the west from Alaska (Moser et al. 1979) and Wyoming (Eversman et
al. 2002). Remarkably, the species has not been reported until now from the
Pacific Northwest, though it is mapped for the Alberta Rockies by Thomson
(1997); the latter specimen has not been seen by us. It is characterized by its
darkly pigmented hypothecium, lack of K+ mauve epihymenial pigments and
3-septate ascospores (see illustrations in Czarnota 2007). It is new to British
Columbia.
SPECIMEN EXAMINED: CANADA. British Columbia. Selkirk Mountains, E of Nakusp,
Kuskanax River drainage, slope below Kimbol Lake, in old-growth Tsuga heterophylla
forest, lignicolous on Thuja plicata wood, 9 Aug 2004, Spribille 15668, Pettitt & Wagner
(hb. Spribille).
Pycnora leucococca (R. Sant.) R. Sant.
This species is known only in the sterile state, but is highly distinctive on
account of its lip-shaped soralia and presence of alectorialic acid. The species
has rarely been reported for North America, but is common in moist areas of
the west, where it was first reported for British Columbia by Moberg (1986,
but unvouchered), Brodo (1991, as an associate of Ochrolechia gowardii) and
Goward & Thor (1992). Brodo (1991) also reported it as an associate of his
Ochrolechia sp. 8 in the Labrador-Québec border region of eastern Canada. It
occurs on a wide array of bark and wood substrates. We report the species here
from southern Nunavut, a substantial range extension halfway between the two
currently known eastern and western ranges in North America; this specimen
too was an incidental collection, an associate of Xylographa opegraphella
New lichen records in North America ... 463
collected in a study by Kershaw in what was then still a part of northwestern
Ontario. We also report it as new to the United States (Montana).
REPRESENTATIVE SPECIMENS EXAMINED (>40 seen): CANADA. Nunavut. East Pen
Island, 56°45°N, 88°45’W, raised beach systems, on drift wood, Jul 1971, Kershaw s.n.
(CANL.-38115, filed under Xylographa opegraphella). U.S.A. Montana. Lincoln Co.,
Yaak Valley, Can Creek, c. 48°44.899’N, 115°38.239'W, 1105 m, corticolous on Abies
lasiocarpa, 7 Nov 2007, Spribille s.n. (hb. Spribille).
Pycnora praestabilis (Nyl.) Hafellner
This species is similar to Pycnora xanthococca but is more robust (in our
material) and has longer conidia (Timdal 1984). A specimen has also been
seen by us from Idaho (GZU) but has apparently been misplaced in that
herbarium.
SPECIMEN EXAMINED: U.S.A. Montana. Ravalli Co., Bitterroot Range, summit of Ingomar
Peak, 46°13°N, 114°25’W, 2590 m, 29 Aug 1982, McCune 12721 (hb. McCune).
Pycnora sorophora (Vain.) Hafellner
Like Pycnora leucococca, this species is often overlooked and rarely reported,
and also as with that species, it is characterized the presence of alectorialic acid
and develops a pinkish colour over time in the herbarium. Unlike P leucococca, it
develops terminal, not marginal soralia and the thallus often quickly completely
disintegrates into soredia (Tonsberg 1992). It is widespread and occurs mostly
on wood although it has been found on both wood and bark in western North
America (Arizona: Timdal 2002; California: Williams & Sillett 2007; Montana:
Hauck & Spribille 2005). We report the species here as new to Alaska, British
Columbia, Colorado, Idaho, Manitoba, Nunavut and Washington.
REPRESENTATIVE SPECIMENS EXAMINED (>50 seen): CANADA. British Columbia.
Thompson Plateau, 20 km NW of Kamloops, Opax Mountain Silvicultural Study Site,
on wood of conifer snag 50°49’N, 120°28’W, Sep 2006, Bjork 13420 (UBC); Manitoba.
5 miles S of Putahow Lake, 59°45’N, 100°43’W, in Picea mariana-Cladina stand on
fallen black spruce, 18 Jun 1973, Wong 1006 (CANL, filed under Xylographa abietina);
Nunavut. East Pen Island, 56°45’N, 88°45’W, raised beach systems, on drift wood,
Jul 1971, Kershaw s.n. (CANL., filed under Xylographa vitiligo, mixed with Pycnora
leucococca); U.S.A. Alaska. Skagway Fjord, west side of Taiya River, 59°32.568’N,
135°20.693’W, 25 m elev., corticolous on Picea sitchensis trunk, 6 Oct 2007, Spribille
24748 (hb. Klondike NHP); Colorado. Clear Creek Co., N face of Squaw Mountain along
Squaw Pass Road, 39°41°30”N, 105°30’W, on wood, 16 Aug 1983, Nash 22134 (ASU, filed
under Xylographa parallela); Idaho. Bonner Co., Cabinet Mountains, University of Idaho
Experimental Forest, Antelope Lake, 48°8’N, 116°10’W, on wood of conifer snag, Bjork
14229 (UBC); Washington. Pend Oreille Co., Frater Lake, 48°39.266’N, 117°29.090°W,
968 m, on snag, 19 Aug 2005, Spribille 17469 (WTU); Spokane Co., Riverside State Park,
near the Bowl & Pitcher overlook, 47°42’N, 117°30’W, wood of Pinus ponderosa log on
talus, Bjork 14135 (UBC).
464 ... Spribille & Bjérk
Pycnora xanthococca (Sommerf.) Hafellner
Pycnora xanthococca differs from the two previous species of Pycnora in
being consistently fertile, but shares with them the characteristic presence of
alectorialic acid in the thallus. In northern Europe, P xanthococca is considered
to be an obligate lignicole (Spribille et al. 2008b), but we have found it in
western North America also on bark of Pseudotsuga menziesii. It has otherwise
been reported from California (Ryan & Nash 1991, Williams & Sillett 2007),
South Dakota (Wetmore 1967), North Dakota (Wetmore 1985) and Wyoming
(Eversman et al. 2002). We report P xanthococca here as new to northwestern
North America.
SPECIMENS EXAMINED: CANADA. British Columbia. Slocan Valley, Perry Ridge trail,
corticolous on Pseudotsuga, 26 Jun 2004, Spribille 15173 (CANL); U.S.A. Idaho. Bonner
Co., Priest River Experimental Forest, behind headquarters, 48°21.30°N, 116°50.19°W,
730 m, lignicolous on Pseudotsuga branches, 24 Jul 2004, Spribille 15342 (BG, hb.
Spribille); Montana. Lincoln Co., northern Salish Mountains, bedrock meadow along
Forest Service Rd. 494, 48°37.224’N, 115°04.037'W, 1491 m, on Pseudotsuga bark in
island of trees in bedrock meadow, 2 Aug 2004, Spribille 15408 & Wagner (hb. Spribille);
same locality, 23 Jul 2006, Spribille 20470 & Abrahamczyk (BG, CANL).
Xylographa trunciseda (Th. Fr.) Minks ex Redinger
This species has been previously reported in North America only from the
Queen Charlotte Islands (Brodo 1995), but is in fact common in inland British
Columbia (e.g., Bunnell et al. 2008) south to northwest Montana and northern
Idaho in humid forests. The species can be easily recognized by the presence
of regularly (American-) football-shaped apothecia produced in chains, with
younger apothecia leading each chain and older apothecia often dying out, and
the presence of the unknown substance referred to by Brodo (1992) as “Xhn-1”
(in TLC producing a LW UV+ blue spot in Rf classes A1/2, B3, C2, yellowish
after charring). The only described species with which it might be confused is
Xylographa hians Willey, which has a nearly identical habit but possesses stictic
acid (Brodo 1992). Xylographa trunciseda is a loyal obligate lignicole wherever
it occurs. It is so far known from Europe and the previous report from British
Columbia; we report it here as new to eastern North America (New Brunswick,
Newfoundland) and the United States (Colorado, Idaho, Montana).
REPRESENTATIVE SPECIMENS EXAMINED (>20 seen): CANADA. New Brunswick. Kent
Co., Kouchibouguac National Park, Rankin’ Brook, 46°50°N, 64°59’W, steep bank to
brook with Tsuga canadensis and Pinus strobus, on dead conifer stump, 23 Aug 1978,
Egger 884 (CANL); Newfoundland. Gros Morne National Park: 3.2 km SW of St. Paul's
(35 km N of Rocky Harbour), in black spruce island with black spruce, balsam fir, birch
and alder, 24 Jun 1981, Wetmore 42781 (CANL, filed under Xylographa vitiligo); U.S.A.
Colorado. Garfield Co. White River Natl. Forest NW of Meadow Lake Campground
on E side of road, Hale, 39°48’N, 107°33W, on wood, 24 Jun 1992, Nash 31936 (ASU);
Idaho. Bonner Co., Canyon Creek Research Natural Area, 48°21.97’N, 116°47.96’W, 790
m, lignicolous on fallen wood, 24 Jul 2004, Spribille 15365 (hb. Spribille); Shoshone
New lichen records in North America ... 465
Co., North Fork Coeur d’Alene River, 47°40’N, 115°59’W, wood of conifer log, 1000 m,
May 2004, Bjérk 8643 (UBC); Montana. Lincoln Co., northern Salish Mountains, Gray
Creek, 48°30.432’N, 114°57.048’W, 1168 m, on rotten log, 9 Aug 2006, Spribille 21821
(hb. Spribille).
Xyloschistes platytropa (Nyl.) Vain.
This species was described from Finland by Nylander (1868) as a Xylographa
and assigned to the new monotypic genus by Vainio (1883), who added new
records from Karelia and Lappland. It has rarely been reported since, e.g., from
Spain (Etayo 1990, as a new variety; also Aragon et al. 2004), Mongolia (Biazrov
et al. 1989) and the Russian Far East (Sikhote-Alin Mountains; Chabanenko
2002). The species is characterized by muriform ascospores 20-30 x 6-8 um,
1 per ascus (rarely 2; Nylander 1868), and immersed thallus forming a whitish
patch on wood. The systematic position of the genus is poorly studied but it is
thought to have affinities to the Ostropales. It is new to North America.
SPECIMEN EXAMINED: CANADA. British Columbia. West Kootenay region, Mount
Revelstoke National Park, Nelson Historic Site, 51°01’N, 118°12’W, 580 m, decorticated
branch of Pinus monticola in rain forest, 2 Aug 2005, Goward 05-242 (UBC).
Acknowledgments
Field work by TS on lignicolous lichens in western Canada was supported in part by
the British Columbia Forest Sciences Program (grant Y062045 to EL. Bunnell) and in
part by the Valhalla Wilderness Society (New Denver, British Columbia). Thanks are
extended to Brian Coppins, Anders Nordin, Goran Thor and Leif Tibell for assistance,
to Zdenék Palice for helping us understand Lecidea symmictella, to Trevor Goward for
allowing us to cite his specimens, to the curators of the herbaria ASU, CANL and FH
and Bruce McCune for loans to TS during his stay in Chicago in March 2008, and to
Thorsten Lumbsch for organizing that stay.
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MYCOTAXON
Volume 105, pp. 469-472 July-September 2008
Coltricia fragilissima, a new record for Brazil
ALLYNE CHRISTINA GOMES-SILVA', LEIF RYVARDEN’
& TATIANA BAPTISTA GIBERTONI!
allynefungi@hotmail.com, leif.ryvarden@bio.uio.no, tbgibertoni@hotmail.com
' Universidade Federal de Pernambuco, Departamento de Micologia
Av. Nelson Chaves s/n, CEP 50760-420, Recife, PE, Brazil
* University of Oslo, Department of Botany
P.O. Box 1045, Blindem, N-0316, Oslo, Norway
Abstract — Coltricia fragilissima is reported from Parque Natural Municipal de Porto
Velho, in the State of Rond6nia, Brazilian Amazonia. This is only the third collection
of this taxon from South America and the first report from Brazil. A description of C.
fragilissima and a key to the six species of Coltricia reported from Brazil are provided.
Key words — fungi, Hymenochaetaceae
Introduction
Coltricia is a cosmopolitan genus described by Gray (1821) characterised by its
stipitate, light-coloured basidiomata, monomitic hyphal system, hyaline to pale
rusty brown, simple septate generative hyphae, and cylindrical to ellipsoidal
basidiospores that are golden yellow to rusty brown and slightly dextrinoid in
Melzer’s reagent (Ryvarden 2004).
Five species of Coltricia have been recorded from Brazil: C. barbata Ryvarden
& de Meijer, C. cinnamomea (Jacq.) Murrill, C. duportii (Pat.) Ryvarden, C.
perennis (L.) Murrill, and C. hamata (Romell) Ryvarden. Some references about
the genus in Brazil are found in the works of Torrend (1926, 1938), Fidalgo
(1968), Ryvarden & de Meijer (2002), and Gibertoni et al. (2004), which
reported or described species collected in North, Northeast and South Brazil.
C. fragilissima was described from French Guyana, and has been reported from
Nigeria and Venezuela (Ryvarden 1982, 2004).
A description of C. fragilissima based on a collection from the Brazilian
Amazonia and a key to the Brazilian species of the genus are presented below.
470 ... Gomes-Silva, Ryvarden & Gibertoni
Material and methods
The Parque Natural Municipal de Porto Velho is located in the state of Rond6nia.
It covers an area of 390.82 1ha, of which 80% is composed of open ombrophilous
forest and the rest by secondary vegetation.
The basidiomata were analysed macro- (shape, colour, hymenial surface) and
micromorphologically (hyphal system, presence/absence and size of sterile
structures and basidiospores). Micromorphological observations were made
from slide preparations in 5% KOH, stained with 1% of aqueous phloxine, and
Melzer’s reagent (Ryvarden 1991). Colour designation followed Watling (1969).
The specimens were deposited in the Herbarium Dr. Ary Tupinamba Penna
Pinheiro (HFSL), Faculdade Sao Lucas, Rondénia, and in the Herbarium Padre
Camille Torrend (URM) of the Universidade Federal de Pernambuco, Brazil.
Taxonomy
Coltricia fragilissima (Mont.) Ryvarden, Nordic J. Bot. 2(1): 78 (1982).
Basidiomata annual, laterally stipitate, pileus circular, infundibiliform, 3.5-3.7
x 2.2-2.4 cm. Stipe short, 3.5 x 1.5 cm, with a basal mycelial disk at the base,
velutinus, fulvous (12). Abhymenial surface glabrous to velutinus, fulvous (12).
Margin entire, involute when dry. Context thin. Hymenial surface poroid, pores
angular, 2-4 mm, fulvous (12). Hyphal structure monomitic; generative hyphae
hyaline, simple-septate, golden brown, 3.5-5 um. Setae absent. Basidia clavate,
15-18 x 4-6 um. Basidiospores hyaline, oblong ellipsoid, smooth, thin-walled,
5-6 (7) x 3-3.5 (4) um.
MATERIAL EXAMINED: BRAZIL. Rondénia: Porto Velho, Parque Natural Municipal de
Porto Velho, XII.2007, A. C. Gomes-Silva 69-70 (URM 78898, URM 78899).
REMARKS: Previous studies of C. fragilissima were based on material collected
in Venezuela and Nigeria (Ryvarden 1982, 2004). The species is characterized
by the short stipe, azonate abhymenial surface, basidiomata of homogeneous
colour, and basidiospores ellipsoid and hyaline (6-7 (8) x 3-3.5 (4) um). C.
barbata differs by the zonate abhymenial surface, longer stipe, and globose
to subglobose basidiospores (5-6 um) (Ryvarden & de Meijer 2002), while
C. cinnamomea has oblong to ellipsoid, golden yellow basidiospores (6.5-8 x
5-6 um) (Ryvarden 2004). The specimens of C. cinnamomea collected in areas
of restinga were initially identified as C. duportii (Rajchenberg & de Meijer
1990, Ryvarden & de Meijer 2002), which differs by having ellipsoid, brown
basidiospores (8-10 x 6-7 um). C. perennis was reported by Ryvarden & de
Meijer (2002), who considered it an exotic species, because it is believed that
it was probably introduced in Brazil along with the plantations of Pinus. C.
hamata is distinguished by its dark, hooked setae. The Brazilian C. fragilissima
Coltricia fragilissima new to Brazil ... 471
differs macromorphologically from the description of Ryvarden (2004) by the
presence of a mycelial disk at the base, and micromorphologically by having
slightly larger basidiospores.
Key to the Brazilian species of Coltricia
farmctac, Ofssetal fivphae present: Weanet Ast setae water ee Let. COMP eleete aw C. hamata
Bmore cae Ol setal NY Diac-dOSCIUEer mr wmer mon ee erm a teaeu/i ac. sence ae ene 2
PM MASICIOSPOTES UDO © [Lk eae eta Oe Meatet PER AR hes wh, cee oe cela ue coe 3
Pipes astCLOSDOLeSTOTIOe! Chlall Ostler erarea ne, ene raeee Meret ee eee me eee ae a
3a. Margin ciliate, 4-9 pores/mm, basidiospores golden yellow to rusty brown,
slobose to’suslobosee aay Meee PRs. Cees ce ee C. barbata
3b. Margin not ciliate, 2-4 pores/mm, basidiospores hyaline,
oblong ellipsOldmaer ee ee etree mer ee eae ea er one C. fragilissima
4a. Abhymenial surface azonate, basidiospores rusty brown .............. C. duportii
apeabhymenial surtace zonate, basidiospores yellowish..-...--.+..--..25-..-+..-- 5
5a. Basidiospores pale yellowish brown, ellipsoid to cylindrical ellipsoid,
ExXOUIC ISD ECIeSaSSOCIALCC Wil gi) smrnean wer ere er eee ee eee C. perennis
5b. Basidiospores golden yellow, oblong to ellipsoid, cosmopolitan species
Picea woes sarc, -Asites ea aoad,-ouid salty: deserhiat dekh. C. cinnamomea
Acknowledgments
The authors would like to thank Ms. Ana Cristina R. Souza, curator of HFSL, for support;
Conselho Nacional de Desenvolvimento Cientifico (CNPq) for the master scholarship
of ACGS, and Dr. Gregory Mueller and Dr. Annarosa Bernnichia for critically reviewing
the manuscript.
Literature cited
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Rain Forest in Northeast Brazil. Synopsis Fungorum 18: 33-43
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Gray SF. 1821. A natural arrangement of British plants. London.
Rajchenberg M, de Meijer AAR. 1990. New and noteworthy polypores from Parana and Sao Paulo
States, Brazil. Mycotaxon 38: 173-185.
Ryvarden L. 1982. Type studies in the Polyporaceae 11. Species described by JFC Montagne, either
alone or with other authors. Nordic Journal of Botany 2 (1): 75-84.
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Fungiflora, Oslo, 363 p.
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Ryvarden L, Gilbertson RL. 1994. European Polypores. Vol. 2. Synopsis Fungorum 6. Fungiflora,
Oslo, 355 p.
Ryvarden L, Johansen I. 1980. A preliminary polypore flora of East Africa. Fungiflora, Oslo, 636 p.
472 ... Gomes-Silva, Ryvarden & Gibertoni
Ryvarden L, de Meijer AAR. 2002. Studies in neotropical polypores 14. New species from the state
of Paranda, Brazil. Synopsis Fungorum 15: 34-69.
Torrend C. 1926. Les polyporacées stipitées du Brésil IV. Brotéria, sér. bot. 22: 5-19.
Torrend C. 1938. As poliporaceas da Bahia e estados limitrofes. Anais da Primeira Reuniao Sul-
Americana de Botanica 2: 325-341.
Watling R. 1969. Colour Identification Chart. Her Majesty's Stationary Office, Edinburgh
(Scotland).
MYCOTAXON
Volume 105, pp. 473-479 July-September 2008
Notes on two boletes with tiny basidioma in tropical China
D. Q. ZHOU' & ZHU L. YANG?
' dqzhou@tnc.org.cn
Kunming University of Science and Technology
Kunming 650093, Yunnan, China
°Correspondant author, fungi@mail.kib.ac.cn
Key Laboratory of Biodiversity and Biogeography, Kunming Institute of Botany
Chinese Academy of Sciences, Kunming 650204, Yunnan, China
Abstract — Two species of boletes with minute basidioma, Boletus coccineinanus and
B. patouillardii, were described and illustrated based on materials made in tropical
Yunnan, southwestern China. Both of them are new to China. Comparisons to other
similar species were made. Boletus minimus, originally described from southeastern
China, is regarded as a synonym of B. patouillardii.
Key words — taxonomy, morphology, Boletales, Basidiomycota
Introduction
During a study of boletes from Yunnan Province, China, we came across two
boletes with minute basidioma. They could easily be overlooked in the field
due to their small sizes. Descriptions and illustrations for the each species are
provided herein, and they are compared with similar species described from
China and Japan.
Materials and methods
Revived tissues were mounted in 5% KOH for microscopic examination. The
abbreviation [n/m/p] shall mean n basidiospores measured from m basidiocarps
of p collections. Dimensions of basidiospores excluding the apiculus are given
using notation of the form (a) b-c (d). The range b-c contains a minimum of
90% of the measured values. Extreme values a and d are given in parentheses.
Q refers to the length/width ratio of basidiospores; Q refers to the average Q of
all basidiospores + sample standard deviation. Herbarium abbreviations follow
Holmgren et al. (1990) but with one exception that is not included in Index
Herbariorum: HKAS—the Herbarium of Cryptogams, Kunming Institute of
Botany, Chinese Academy of Sciences.
474 ... Zhou & Yang
Taxonomy
Boletus coccineinanus Corner,
Boletus in Malaysia (Singapore): 152, Fig. 47, Pl. 11/2 (1972). Figs. 1-4
BASIDIOMATA (Fig. 1) very small. PILEus 10-12 mm in diam., convex to plano-
convex; surface dry, red to vinaceous red, often cracked into minute squamules;
context white to whitish, unchanging. HyMENOPHORE poroid, carneous,
unchanging when bruised, slightly depressed around apex of stipe; pores 2-
3/mm, but becoming somewhat larger towards apex of stipe; color of tubes
unknown. STIPE 12-20 x 2-3 mm, subequal, often curved near apex or base,
smooth, without nets, sometimes with longitudinal striations, nearly glabrous
or with very small fibrillose squamules, dry, mostly concolorous with pileus,
whitish at apex, with ochre to yellow mycelium at base; context yellowish but
ochreous yellow at base, unchanging. ANNULUs absent.
TUBE TRAMA boletoid with deep-colored mediostratum and light-colored
lateral strata. BAstp1a 25-30 x 8-10 um, clavate, hyaline, 4-spored, no clamps
observed on basal septa. BAstDIosporEs (Fig. 2) [35/2/1] (8) 8.5-10.5 (11) |
x 4-5 um, Q = (1.55) 1.88-2.33 (2.44) (Q = 2.0 + 0.17), yellowish in KOH,
slightly thick-walled (< 0.5 um), smooth, boletoid to subamygdaliform in side
view and usually without a suprahilar depression, nearly ellipsoid in ventral
view. CHEILOCYSTIDIA (Fig. 3) 30-40 x 6-8 um, subfusiform to narrowly
clavate, thin-walled, nearly colorless and hyaline. PLEUROCyYsTIDIA similar to
cheilocystidia in form and size, but rare. PILEIPELLIS (Fig. 4) an epithelioid
trichoderm composed of more or less vertically arranged, moniliform hyphae
with ovoid, subglobose to ellipsoid, often thick-walled (up to 1.5 um thick),
reddish brown vacuolar pigmented cells (10-30 x 8-15 um) 2-4 in chains.
Camps absent in all tissues.
HABIT, HABITAT, DISTRIBUTION AND SEASON — Single or in groups, on soil
in tropical forest; Tropical China, Indonesia (Corner 1974), Malaysia (Corner
1972), and Thailand (Watling 2001); August-November.
SPECIMEN EXAMINED — CHINA. YUNNAN PROVINCE: Longchuan County, along a
roadside near Nongxianhe, alt. 1200 m, 25.VII.2003, H. Luo 62 (HKAS 43601).
COMMENTS—Boletus coccineinanus is well characterized by its very small
basidioma with a red pileus, a red stipe with an ochre basal mycelium, a pink
to pinkish hymenophore, relatively subamygdaliform basidiospores and an
epithelioid trichoderm of the pileipellis with thick-walled cells. Corner (1972)
put this species in the alliance around B. nanus of Boletus subgen. Tylopilus.
Because molecular phylogenetic studies of many species of boletes including
B. coccineinanus are still unavailable, we prefer to keep it in the genus Boletus
temporarily. The characters observed from the Chinese material agrees
relatively well with the protologue of B. coccineinanus. In the protologue the
Boletus coccineinanus & B. patouillardii new to China ... 475
20 pm
IN
Figs. 1-4: Boletus coccineianus (HKAS 43601).
1. Basidiomata; 2. Basidiospores; 3. Cheilocystidia; 4. Pileipellis.
cheilocystidia were described as clavate with pink walls. The pink wall probably
can only be observed in water with fresh material. In the Chinese material the
pink color of the cell wall couldn't be discernible. This species has not been
reported from China before (see Bi et al. 1994, 1997; Zang 2006).
Boletus patouillardii Singer, Am. Middl. Nat. 37: 55 (1947). Figs. 5-9
= Porolaschia bicolor Pat., Bull. Soc. Mycol. France 39: 54 (1923).
= Boletus minimus M. Zang & N.L. Huang, Acta Bot. Yunnanica 24: 723, fig. 1/1-6 (2002).
BASIDIOMATA (Fig. 5) very small. PrLEus 8-17 mm in diam., convex to plano-
convex; surface dry, dark red to orange red when fresh, becoming dark brown
when dried, nearly glabrous to finely tomentose; context yellowish, unchanging.
HyYMENOPHORE poroid, yellowish to yellow at first, but becoming pale yellow-
brown when mature, slightly depressed around apex of stipe; pores 2(3)/mm,
but becoming larger towards apex of stipe and 1/mm, and often forming
lamella-like “teeth” adnate to apex of stipe and, thus, hymenophore somewhat
appearing decurrent. STIPE 10-20 x 1-3 mm, subcylindrical, smooth, without
nets or striations, nearly glabrous or with fine fibrillose squamules, dry,
vinaceous, with yellow to yellowish mycelium at base. ANNULUS absent.
TuBE TRAMA boletoid with deep-colored mediostratum and light-colored
lateral strata. BAstD1A 25-30 x 7-10 um, clavate, hyaline, 4-spored, no clamps
observed on basal septa. BAstDIOSsPoRES (Figs. 6-7) [63/3/2] (6.5) 7-8.5 (9) x
4—5 um, Q = (1.56) 1.63-1.90 (2.0) (Q = 1.78 + 0.11), ochreous to yellowish,
slightly thick-walled (< 0.5 um), smooth, ellipsoid to subamygdaliform in side
476 ... Zhou & Yang
Figs. 5-8: Boletus patouillardii.
5. Basidiomata (holotype); 6. Basidiospores (holotype);
7. Basidiospores (HKAS 43602); 8. Pileipellis (HKAS 43602).
view and usually without a suprahilar depression, ellipsoid in ventral view.
CHEILOCYSTIDIA 30-50 x 5-8 um, subfusiform to fusiform, nearly colorless
and hyaline, thin-walled. PLEUROCyYsTIDIA similar to cheilocystidia in size and
form, but rare. PILEIPELLIS (Figs. 8-9) an epithelioid trichoderm composed of
more or less vertically arranged, moniliform hyphal elements 2-4 in chains,
10-40 x 7-16 um, ellipsoid, ovoid, subglobose, thin-walled with yellowish,
yellow-brown to brownish vacuolar pigment; terminal ones broadly ellipsoid
to subfusiform or subcylindrical, with round to obtuse apices. CLAMPs absent
in all tissues.
HABIT, HABITAT, DISTRIBUTION AND SEASON — Single or in groups, on soil in
forest; Cambodia, and tropical and subtropical China; July-August.
SPECIMENS EXAMINED — CAMBODIA. KOMPONG CHHNANG: Forest Reserve, 1921,
M. Petelot 222 (HOLOTYPE of Porolaschia bicolor and Boletus patouillardii, FH-Pat.
Herb.3644). CHINA. YUNNAN PROVINCE: Longchuan County, along a roadside near
Nongxianhe, alt. 1200 m, 25.VIII.2003, H. Luo 63 (HKAS 43602); FUJIAN PROVINCE:
Yongchun County, Niumulin Nature Reserve, 3.VII.2001, N.L. Huang 703 (HKAS
39514, HOLOTYPE of Boletus minimus). ;
COMMENTS—Boletus patouillardii is well characterized by its very small
basidioma with a red pileus, and a red stipe with a yellow basal mycelium, a
yellowish to yellow hymenophore unchanging in color when injured, relatively
short ellipsoid to subamygdaliform basidiospores and an epithelioid trichoderm
of the pileipellis.
Boletus coccineinanus e& B. patouillardii new to China ... 477
Fig. 9: Boletus patouillardii.
Pileipellis (holotype of B. minimus).
The holotype specimen did not rehydrate well, thus microscopic studies were
limited. Cystidia and pileipellis were not recovered, and basidiospores were
[42/2/1] (6.5) 7-9 x 4-5 um, Q = (1.56) 1.63-1.89 (2.0) (Q = 1.75 + 0.11).
The name Porolaschia bicolor was published by Patouillard (1923), based on
the same type specimen. Singer was aware that Patouillard had examined the
specimen and named it as a species of Porolaschia, but mistakenly thought that
this was merely a herbarium name (Singer 1945, 1947). Porolaschia bicolor
cannot be recombined in Boletus, because the epithet is already occupied by
Boletus bicolor Peck; thus Singer's epithet is the earliest available in Boletus for
this taxon. Although published as a “sp. nov.’, Boletus patouillardii should be
regarded as a nom. nov. for its homotypic synonym, Porolaschia bicolor. This
species was known only from its type collection.
Boletus minimus is apparently conspecific with B. patouillardii due to its
minute basidioma with a red pileus, a red stipe and a yellow hymenophore.
Although the base of the stipe was described as “whitish’, reexamination of
the holotype of B. minimus showed that the mycelium at the base of the stipe
is yellowish. The only basidioma available for our study is unfortunately not
mature. In the protologue, the basidiospores of B. minimus were described
as “10.4-14 x 4-5.2 um’ longer than those of B. patouillardii. However, the
line-drawings (Zang & Huang 2002, Fig. 1/4; Zang 2006, Fig. 36/4) showed
that the size and the form of the spores are just within the ranges of those of
B. patouillardii. Furthermore, the pileipellis of the holotype of B. minimus (Fig.
9) is very similar to that of B. patouillardii. Thus, B. minimus should be regarded
as a synonym of B. patouillardii.
Another species with small basidiomata, Xerocomus tengii M. Zang et al.,
was described from southern subtropical China (Zang et al. 2002). Restudy
478 ... Zhou & Yang
of the type of X. tengii (HKAS 39594) revealed that the surface of the pileus
is rugulose and in some areas even distinctly reticulate, the basidiospores are
10-14 x 4-5 um, and the pileipellis is an ixotrichoderm consisting of more
or less vertically arranged, frequently septate filamentous hyphae, very similar
to that of Aureoboletus thibetanus (Pat.) Hongo & Nagas. (Yang et al. 2003).
However, cystidia with refractive substance on its surface, very characteristic
for A. thibetanus, were not observed from the holotype.
Boletus aokii Hongo, originally described from Japan, looks like B.
patouillardii, but differs from the latter by its basidioma becoming blue on
injury, longer basidiospores, and subventricose, apically tapering terminal cells
in the pileipellis (Hongo 1984).
Acknowledgments
This study was supported by grants from the National Natural Science Foundation of
China (No. 30525002) and the Knowledge Innovation Program of the Chinese Academy
of Sciences (KSCX2-YW-G-025). The authors are very grateful to Prof. Dr. D. H. Pfister,
Farlow Herbarium of Harvard University, USA for allowing the authors access to the
specimens in the Herbarium, and using his laboratory to study the collections, to
Prof. Dr. T. H. Li, Guangdong Institute of Microbiology, China, and Mr. E. Nagasawa,
the Tottori Mycological Institute, Japan, for serving as reviewers. Dr. H. Luo, Yunnan
University, China, is acknowledged for providing fungal materials.
Literature cited
Bi ZS, Li TH, Zhang WM, Song B. 1997. A preliminary agaric flora of Hainan Province. Guangdong
Higher Education Press: Guangzhou (China). 388 pp.
Bi ZS, Zheng GY, Li TH. 1994. Macrofungus flora of Guangdong Province. Guangdong Science and
Technology Press: Guangzhou (China). 879 pp.
Corner EJH. 1972. Boletus in Malaysia. Government Printer: Singapore. 263 pp.
Corner EJH. 1974. Boletus and Phylloporus in Malaysia: further notes and descriptions. The
Gardens’ Bulletin Singapore 27:1-16.
Holmgren PK, Holmgren NH, Barnett LC. 1990. Index herbariorum. Part I: herbaria of the world.
8th edition. New York Botanical Garden: New York (USA). 693 pp.
Hongo T. 1984. Materials for the fungus flora of Japan (35). Transactions of the Mycological Society
of Japan 25: 281-285.
Patouillard N. 1923. Herborisations mycologiques au Cambodge. Bulletin dela Societé Mycologique
de France 39: 46-58.
Singer R. 1945. The Laschia-complex (Basidiomycetes). Lloydia 8: 170-230.
Singer R. 1947. The Boletoideae of Florida with notes on extralimital species III. The American
Midland Naturalist 37: 1-135.
Watling R. 2001. The relationships and possible distributional patterns of boletes in south-east
Asia. Mycological Research 105: 1440-1448.
Yang ZL, Wang XH, Binder M. 2003. A study of the holotype and additional materials of Boletus
thibetanus. Mycotaxon 86: 283-290.
Boletus coccineinanus & B. patouillardii new to China ... 479
Zang M. 2006. Flora fungorum sinicorum. Vol. 22 Boletaceae (I). Science Press: Beijing (China).
215 pp.
Zang M, Huang NL. 2002. A new species of the genus Boletus from China, Boletus minimus. Acta
Botanica Yunnanica 24: 723-724.
Zang M, Lin JT, Huang NL. 2002. Xerocomus tengii, a new Xerocomus species from China.
Mycosystema 21: 480-482.
MYCOTAXON
Volume 105, pp. 481-488 July-September 2008
The generic name Leratiomyces (Agaricales) once again
ScoTT A. REDHEAD’ & JOHN MCNEILL?’
redheads@agr.gc.ca
' National Mycological Herbarium, Eastern Cereal & Oilseed Research Centre C.E.E,
Agriculture & Agri-Food Canada
Ottawa, Ontario, Canada, K1A 0C6
jmcneill@rbge.ac.uk
? Royal Botanic Garden
Edinburgh, EH3 5LR, Scotland, U.K.
Abstract — Leratiomyces was not validly published in 1998; rather it was, in fact,
unintentionally validated in 2008 as a nomen novum for Le-Ratia. The history of these
names is detailed and linked to “Le Ratia,” which is not a validly published name.
Le-Ratia, which is validly published but illegitimate, is lectotypified, thereby typifying
Leratiomyces; the new combination Leratiomyces similis is proposed for its type.
Key words — New Caledonia, nomenclature, Orthotrichaceae, secotioid, Stropharia
Introduction
The generic name Leratiomyces was adopted for a phylogenetically defined clade
of agarics and secotioid gasteromycetes in a recent publication in Mycotaxon
(Bridge et al. 2008) following earlier analyses by Binder et al. (1997), Bresinsky
& Binder (1998), Moncalvo et al. (2002), and Walther et al. (2005). However,
the cited type, “L. similis” (Bridge et al. 2008), was based on a name that was
not validly published. This contradiction led us to investigate the nomenclature
behind the name Leratiomyces.
History
The name “Leratiomyces” was proposed by Bresinsky & Binder (1998) as
a ‘nomen novum’ for “Le Ratia” (Patouillard 1907) that was never validly
published. New replacement names, nomina nova, can only be proposed for
validly published names (Art. 7.3 & 33.4); ifa name is not validly published
© Her Majesty the Queen in Right of Canada, as represented by the Minister of Agriculture and
Agri-Food Canada
482 ... Redhead & McNeill
it has no status (Art. 12.1). Hence use of “names” in the International Code of
Botanical Nomenclature (McNeill et al. 2006) refers only to validly published
names (Art. 6.3). “Le Ratia” Pat. was not validly published (Art. 20.3 & Ex. 7)
because it consists of a two part “generic” name that, unlike species epithets
that can be corrected (Art. 23.1, 60.9 ), cannot be “corrected” by later authors
by uniting or hyphenating while retaining the original authorship and date
and place of publication for priority purposes. The name Le-Ratia (with the
hyphen) was validly published (Art. 20.3) by Saccardo & Trotter (1912) along
with the binomial Le-Ratia similis for the original taxon detailed by Patouillard
(1907) as “Le Ratia similis’. Additionally, they validated the name Le-Ratia
smaragdina for a species circumscribed in a second paper by Patouillard (1909)
that he called “Le Ratia smaragdina” - also not a validly published name.
Species names, such as those put forward by Patouillard (1907, 1909), cannot
be validly published when the epithet is combined with a purported generic
name that itself is not validly published (Art. 43.1). Authorship for Le-Ratia
must be attributed to Saccardo & Trotter (1912), not Patouillard, and it dates
from 1912, not 1907 or 1909.
Prior to validation of Le-Ratia Sacc. & Trotter in 1912, an earlier homonym,
Leratia Broth. & Paris (Brotherus 1909a,b), was published for a moss in
honour of the same collector, Auguste-Joseph Le Rat, who sent materials from
New Caledonia (cf. Farr et al. 1979). The two names Le-Ratia and Leratia are
interpreted as homonyms even though differing in one being hyphenated. For
example, Brotherus & P. Sydow (Sydow 1910) incorrectly believed that the moss
genus name Leratia was a later illegitimate homonym of “Le Ratia Pat’, leading
them to propose unnecessarily yet another invalid, two part, replacement
generic name “Le Ratiella” for the moss. Leratia continues to be in current
use for mosses (Crum 1987, Goffinet et al. 1998, 2004). Additional support for
treating the two generic names as homonyms comes from Wakefield's (1922)
discussion of both of Patouillard’s species’ specimens, labeling them “LeRatia
similis” and “LeRatia smaragdina’. Interpreted as a later homonym of Leratia,
the fungal Le-Ratia is illegitimate (Art. 53.1 & 53.3), which is one of the reasons
why bryologists continue to use the generic name Leratia. However, the two
species names, Le-Ratia similis and Le-Ratia smaragdinus, dating from 1912
(not 1907 or 1909 respectively) are legitimate (Art. 55.1).
The reasons why “Leratiomyces” (Bresinsky & Binder 1998) was not validly
published, either as the name of a new genus or as a nomen novum for the -
validly published but illegitimate Le-Ratia (Saccardo & Trotter 1912), must
be considered. Art. 33.4 dictates that after 1952 a full and direct reference be
given to the author and place of publication of the replaced synonym to validate
nomina nova. Unfortunately, no reference was made to Saccardo & Trotter
Leratiomyces lectotypified ... 483
(1912) in the 1998 publication. Art. 33.5 allows for correction of bibliographic
citations with regard to Art. 33.4. The definition of correctable citation situations
covered by Art. 33.5 is governed by Art. 33.7, wherein only four case scenarios
are allowed: a) the replaced synonym was validated earlier than the cited source
(this is not the case—see below); b) the incorrect citation resulted from a shift
in starting point dates (this does not apply here); c) the intended generic name
has a basionym (it does not have a basionym); and d) when it would otherwise
be the validly published name of a new taxon (discussed below).
Bresinsky & Binder listed as “Ungiiltige Namen” three differently spelled
names that were neither validly published nor identified by a full and direct
reference: “Le Ratia Pat. 1907. - Le-Ratia, Heim 1957.- Leratia, Smith & Singer
1959, Ainsworth 1961” Hence the fundamental criteria outlined in Art. 33.7(a)
were not met. Heim appears not to have published in 1957, but did so in 1951,
1967 and 1968a,b; only in the 1951 paper did he use the hyphenated spelling,
but did so inconsistently, so “1957” may be a typographical error for “1951”.
Smith & Singer (1959: 216) incidentally discuss “Leratia Pat’, spelled as such,
in reference to both Patouillard’s and Heim’s publications. Ainsworth’s (1961)
publication was the Dictionary of Fungi.
Finally, it must be asked whether the requirements for valid publication
of a name of a new taxon were met (Art. 33.7(d)) when “Leratiomyces” was
proposed by Bresinsky & Binder (1998). This would require provision of a
Latin diagnosis or description (Art. 36.1) and designation of a type (Art. 37.1).
The authors supplied a Latin description but they designated as nomenclatural
type “Le Ratia similis Pat. (siehe Leratiomyces similis)”. For “Leratiomyces
similis (Pat.) Bresinsky et Binder” the basionym was given as “Le Ratia similis
Patouillard, Bull. Soc. Mycol. de France 23, 52, 1907.” “Le Ratia similis Pat.” is
not a validly published name and therefore citation of it cannot be construed
as effectively citing a type because it is not a named species (Art. 37.3). It may
be further asked if the binomial “Leratiomyces similis” were unintentionally
validated as the name of a new taxon by meeting all such requirements. The
reference to Patouillard (1907) leads to a Latin description of the taxon, but
no type was designated for the name in 1998 and therefore the binomial
“Leratiomyces similis” is not validly published. Designation of a type for a
proposed generic name via citation of two names that were never validated is
not an effective designation of a type under Art. 10.1 for the proposed generic
name “Leratiomyces” (Art. 33.4), which itself must therefore be treated as not
validly published. The requirements of Art. 33.7(d) were not met.
Bresinsky & Binder (1998) had intended to include four species and had
proposed “new combinations” based upon “Le Ratia atrovirens Heim’, “Le
Ratia coccinea Massee & Wakefield”, and “Le Ratia smaragdinus Patouillard’,
484 ... Redhead & McNeill
in addition to “Le Ratia similis”; none of these is a validly published name.
None of the combinations is validly published because neither the generic
name “Leratiomyces” nor any one of the basionyms is validly published (Art.
43.1). Additionally, their reference to “Le Ratia atrovirens Heim” referred to a
publication (Heim 1968a) that lacked a Latin description or diagnosis (supplied
subsequently in Heim (1968b)).
Leratiomyces was more recently adopted by Bridge et al. (2008), who proposed
six additional new species combinations and one varietal combination. Although
these authors were evidently unaware of the fact that “Leratiomyces Bresinsky
& Manfr. Binder” was not validly published, they did in fact cite, along with
“Le Ratia Pat.” (acknowledged as a “nom. inval.”), Le-Ratia Pat. ex Sacc. & Trotter
as a nomenclatural (homotypic) synonym of Leratiomyces along with a direct
and full citation to its place of publication in their synonymy for Leratiomyces.
Although they listed the type under two designations, “L[eratiomyces] similis
(Pat.) Bresinsky & Manfr. Binder” and “Le Ratia similis Pat.” neither of which
is a validly published name and did not mention Le-Ratia similis Pat. ex Sacc. &
Trotter, it is not required under Art. 37.1 that a name published as a replacement
for a validly published (but illegitimate) name include citation of the type. Nor
was it necessary in 1912 for Saccardo & Trotter to cite a type. Consequently
Bridge et al., albeit inadvertently, have validly published Leratiomyces, which
must therefore be attributed as Leratiomyces Bresinsky & Manfr. Binder ex
Bridge et al., Mycotaxon 103: 115. 2008.
Despite the validation of the generic name, the binomial “Leratiomyces
similis” for the type of the generic name has yet to be validly published. The
authors, unaware of the fact that Bresinsky & Binder had not validly published
Leratiomyces, did not mention Le-Ratia similis Pat. ex Sacc. & Trotter, far less
give “a full and direct reference ... to its author and place of valid publication”
(Art. 33.4), as would be necessary for their use of “L. similis” to serve as valid
publication of the name. However, the seven proposed combinations of other
species and varieties of Leratiomyces later in the paper are validly published.
One other generic name, “Stropholoma’, was proposed by Balletto (1989) for
species now placed in the “Leratiomyces” clade. However, Balletto intended
to elevate to generic level Naematoloma sect. Stropholoma Singer (1948), but
failed to provide a full and direct reference to the place of publication of the
basionym. Consequently, “Stropholoma” is not a validly published generic
name (Art. 33.4), as noted by Bridge et al. (2008).
No validly published names exist for the two fungal species labeled “Le Ratia
atrovirens’ (Heim 1968a,b) or “Leratiomyces atrovirens” (Bresinsky & Binder
1998) and “Le Ratia coccinea” (Wakefield et al. 1916, Wakefield 1920) or
Leratiomyces lectotypified ... 485
“Leratiomyces coccinea” (Bresinsky & Binder 1998). ‘These will need to be
described as new taxa in appropriate genera if and when names are needed.
We hesitate to validate them with an illegitimate generic name, Le-Ratia, or a
taxonomically inappropriate genus, Leratiomyces. The only validly published
name for the remaining taxon also excluded from Leratiomyces by Bridge et al.
(2008), is Le-Ratia smaragdinus Pat. ex Sacc. & Trotter.
With regard to “Le Ratia coccinea’, we note that the publication of new Latin
descriptions for several fungi by Wakefield et al. (1916), which were mentioned
in a footnote by an editor (H. Schinz) in the publication by Wakefield (1920),
is generally overlooked because Latin descriptions are again provided in 1920.
Apparently the manuscript for the 1920 publication was available in 1914 but
publication was delayed because of World War I [editor's note in Wakefield et al.
1916], and Schinz wished to ensure priority of authorship for the mycologists
for the new taxa despite the delay.
Leratiomyces as currently recognized occurs in both the Northern and Southern
Hemispheres, encompasses both lamellate and secotioid forms, and inhabits
both forested as well as rural and urban habitats. The name Leratiomyces has
already been adopted for agarics by one author (Kibby 2008).
We offer the following nomenclatural synopsis and new combination as a
guide to the names. For more complete taxonomic synonymy, see Bridge et.
al. (2008).
Nomenclature
Leratiomyces Bresinsky & Manfr. Binder ex Bridge, Spooner, Beever & D.-C. Park,
Mycotaxon 103: 115 (2008)
= Le-Ratia Pat. ex Sacc. & Trotter, Syll. fung. 21: 468 (1912) nom. illegit., non Leratia
Broth. & Paris in Brotherus (1909a, b) (Art. 53.1 & 53.3)
= “Le Ratia Pat, Bull. Soc. Mycol. France 23: 52 (1907) non rite publ. (Art. 20.3)
= “Leratiomyces Bresinsky & Manfr. Binder’, Z. Mykol. 64(1): 80 (1998) non rite publ.
(Art. 33.4)
Lectotypus (designated here): Leratiomyces similis (Pat. ex Sacc. & Trotter)
Bresinsky & Manfr. Binder ex Redhead & McNeill, comb. nov.
MycoBANkK MB 512293
BastonyM: Le-Ratia similis Pat. ex Sacc. & Trotter, Syll. Fung. 21: 468 (1912).
= “Le Ratia similis Pat’, Bull. Soc. Mycol. Fr. 23: 52 (1907) non rite publ.(Art.
43.1)
= “Leratiomyces similis (Pat.) Bresinsky & Manfr. Binder’, Z. Mykol. 64(1): 80
(1998) non rite publ. (Art. 43.1)
486 ... Redhead & McNeill
Additional included species:
Leratiomyces ceres (Cooke & Massee) Spooner & Bridge
in Bridge et al., Mycotaxon 103: 116 (2008)
Leratiomyces cucullatus (Shope & Seaver) Beever & D.-C. Park
in Bridge et al. Mycotaxon 103: 116 (2008)
Leratiomyces erythrocephalus (Tul. & C. Tul.) Beever & D.-C. Park
in Bridge et al. Mycotaxon 103: 116 (2008)
Leratiomyces magnivelaris (Peck) Bridge & Spooner
in Bridge et al., Mycotaxon 103: 116 (2008)
Leratiomyces percevalii (Berk. & Broome) Bridge & Spooner
in Bridge et al., Mycotaxon 103: 116 (2008)
Leratiomyces squamosus (Pers.) Bridge & Spooner
in Bridge et al., Mycotaxon 103: 116 (2008)
Leratiomyces squamosus var. thraustus (Schulzer ex Kalchbr.) Bridge & Spooner
in Bridge et al., Mycotaxon 103: 117 (2008)
Excluded species with a validly published name
(see Bridge et al. 2008):
Le-Ratia smaragdinus Pat. ex Sacc. & Trotter, Syll. fung. 21: 468 (1912)
= “Le Ratia smaragdinus Pat’, Bull. Soc. Mycol. Fr. 25: 133 (1909) non rite publ.
(Art. 43.1)
= “Leratiomyces smaragdinus (Pat.) Bresinsky & Manfr. Binder’, Z. Mykol. 64(1): 80
(1998) non rite publ.(Art. 43.1)
Excluded species lacking a validly published name
(see Bridge et al. 2008):
“Le Ratia atrovirens R. Heim’, Rev. Mycol. (Paris) 33: 143 (1968) non rite publ.
(Art. 36.1, 43.1)
= “Le Ratia atrovirens R. Heim’, Rev. Mycol. (Paris) 33: 212 (1968) non rite publ.
(Art. 43.1)
= “Leratiomyces atrovirens (R. Heim) Bresinsky & Manfr. Binder’, Z. Mykol. 64(1): 80
(1998) non rite publ.(Art. 33.4, 36.1, 43.1)
Unclassified enigmatic taxon lacking a validly published name:
“Le Ratia coccinea Massee & Wakef?’ in Wakefield et al., Vierteljahrsschr. Naturf.
Ges. Ziirich 61: 631 (1916) non rite publ.(Art. 43.1)
= “Le Ratia coccinea Massee & Wakef.” in Wakefield, Nova Caledonia, Bot. (ed. F.
Sarasin, J Roux) 1(2): 104 (1920) non rite publ.(Art. 43.1)
= “Leratiomyces coccinea (Massee & Wakef.) Bresinsky & Manfr. Binder’, Z. Mykol.
64(1): 80 (1998) non rite publ.(Art. 43.1)
|
Leratiomyces lectotypified ... 487
Acknowledgements
We thank Shaun Pennycook for supplying references and alerting us to literature and
both him and Paul Kirk for presubmission reviews.
Literature cited
Balletto C. 1989. Problemi di nomenclatura — XV. Micol. Ital. 18(1): 33-37.
Binder M, Besl H, Bresinsky A. 1997. Agaricales oder Boletales? Molekularbiologische Befunde zur
Zuordnung einiger umstrittener Taxa. Z. Mykol. 63: 189-196.
Bresinsky A, Binder M. 1998. Leratiomyces nom. nov. fir eine bislang nicht giiltig beschriebene
Gattung der Strophariaceae (Agaricales) aus Neukaledonien. Z. Mykol. 64: 79-82.
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NOMENCLATURAL NOVELTIES PROPOSED IN MYCOTAXON 105
Aecidium haqii Sultan, Khalid & Mukhtar, p. 25
Aecidium pakistanicum Sultan, I.U. Haq, Khalid & Mukhtar, p. 24
Amanita savannae Tulloss & Franco-Mol., p. 318
Ambispora brasiliensis B.T. Goto, L.C. Maia & Oehl, p. 13
Arthonia hawksworthii Halici, p. 94
Balaniopsis triangularis D.W. Li & W.B. Kendr., p. 106
Boletellus piakaii T.W. Henkel & Fulgenzi, p. 388
Bulbothrix megapotamica Canéz & Marcelli, p. 225
Cruentomycena R.H. Petersen, Kovalenko & O. Morozova, p. 123
Cruentomycena kedrovayae R.H. Petersen, Kovalenko & O. Morozova, p. 127
Cruentomycena viscidocruenta (Cleland) R.H. Petersen & Kovalenko, p. 123
Dactylellina daliensis Hong Yan Su, p. 314
Endoperplexa tortola P. Roberts, p. 140
Entoloma alnicola Noordel. & Polemis, p. 303
Entoloma leuconitens Noordel. & Polemis, p. 302
Exobasidium rhododendri-nivalis Zhen Ying Li & L. Guo, p. 331
Exobasidium pyroloides Zhen Ying Li & L. Guo, p. 332
Fomitiporia ellipsoidea B.K. Cui & Y.C. Dai, p. 344
Ganoderma vivianimercedianum Torres- Torres, p. 448
Gloeoporus dichrous var. niger (Ryvarden) TomSovsky & Ryvarden, p. 173
Gymnopus inexpectatus Consiglio, Vizzini, Antonin & Contu, p. 45
Heterochaete inconspicua P. Roberts, p. 142
Heteroconium avilae R.F. Castaneda, Iturr., Heredia & Minter, p. 176
Inocephalus dragonosporus (Singer) T.J. Baroni & Largent, p. 187
Isaria locusticola Z.Q. Liang, X.Y. He & Y.F. Han, p. 31
Leratiomyces similis (Pat. ex Sacc. & Trotter) Bresinsky & Manfr. Binder
ex Redhead & McNeill, p. 85
Llimoniella muralicola Halici, p. 204
Morchella tomentosa M. Kuo, p. 441
Myelochroa supraflava Canéz & Marcelli, p.227
Parmotrema aberrans (Vain.) Canéz & Marcelli, p. 231
Parmotrema eliasaroanum Benatti, Marcelli & Elix, p. 236
Parmotrema graniticum Benatti, Marcelli & Elix, p. 239
Parmotrema milanezii Marcelli, Benatti & Elix, p. 244
Perenniporia minor Y.C. Dai & H.X. Xiong, p. 60
Phacellium annonae-cherimoliae U. Braun, p. 209
489
490
Phellinus mori Y.C. Dai & B.K. Cui, p. 54
Plasmopara domingensis (Cif.) YJ. Choi & H.D. Shin, p. 194
Pluteus bii Minnis & Sundb., p. 166
Polysporina arenacea (H. Magn.) K. Knudsen & Kocourk., p. 157
Polysporina subfuscescens (Nyl.) K. Knudsen & Kocourk., p. 151
Protomerulius efibulatus Y.C. Dai & Y.L. Wei, p. 350
Pseudocercospora annonarum (Petr. & Cif.) U. Braun & Crous, p. 212
Pseudocercospora asiminae (Ellis & Morgan) U. Braun & Crous, p. 221
Pseudocercospora oblecta (Syd.) Crous & U. Braun, p. 222
Pseudocercospora passiflorae-setaceae A.C. Dianese, A.M. Costa & Dianese, p. 2
Pyriculariopsis formosa Rob. Fernandez, R.F. Castaneda & Iturr., p. 338
Radiigera tropica Orihara & T. Kasuya, p. 112
Repetobasidiopsis Dhingra & Avneet P. Singh, , p. 421
Repetobasidiopsis grandisporus Dhingra & Avneet P. Singh, p. 422
Sarcogyne lapponica (Ach. ex Schaer.) K. Knudsen & Kocourk., p. 160
Scutellospora striata Cuenca & R.A. Herrera, p. 81
Stropharia trinitensis (Dennis) Cortez, p. 10
Tetraploa circinata Pratibha & Bhat, p. 423
Trimitiella Dhingra, p. 422
Trimitiella indica Dhingra, p. 422
Vamsapriya mahabaleshwarensis Pratibha & Bhat, p. 426
Wardomycopsis litoralis Silvera, Gené, Cano & Guarro, p. 197
Xanthoparmelia conglomerata Canéz & Marcelli, p. 228
49]
AUTHOR INDEX— VOLUME ONE HUNDRED FIVE
Afshan, N.S., A.N. Khalid & A.R. Niazi. New records and distribution of rust fungi
from Pakistan. 105: 257-267. 2008.
Aime, M. Catherine, see Largent & al.
Antonin, Vladimir, see Vizzini & al.
Aptroot, André, see Yazici & Aptroot
Baroni, Timothy J., see Largent & al.
Baseia, I.G., see Gurgel & al.
Batzer, Jean C., see Zhai & al.
Benatti, Michel N., see Marcelli & al.
Bhat, D.J., see Pratibha & Bhat
Bjork, Curtis R., see Spribille & Bjork
Boccardo, Fabrizio, see Zotti & al.
Braun, Uwe & Pedro Crous. Cercosporoid hyphomycetes on hosts of the Annonaceae:
Cercospora annonaceae and Isariopsis annonarum revisited. 105: 207-224. 2008.
Cai, Shouping, see Liang & al.
Candan, Mehmet & Aysen Ozdemir Tiirk. Lichens of Malatya, Elazig, Adryaman
Provinces of Turkey. 105: 19-22. 2008.
Canéz, Luciana S., see Marcelli & Canez
Cano, Josep, see Silvera-Simon & al.
Castafieda Ruiz, Rafael E., see Iturriaga & al.
Castafieda Ruiz, Rafael F., Teresa Iturriaga, Gabriela Heredia Abarca, David W. Minter,
Josepa Gené, Marc Stadler, Masatoshi Saikawa & Carolina Silvera-Simon. Notes
on Heteroconium and a new species from Venezuela. 105: 175-184. 2008.
Choi, Young-Joon & Hyeon-Dong Shin. Reclassification of Bremia domingensis to the
genus Plasmopara as P. domingensis comb. nov. 105: 190-194. 2008.
Cifuentes, Joaquin, see Rosique-Gil & al.
Cifuentes, Joaquin, see Sierra & al.
Consiglio, Giovanni, see Vizzini & al.
Contu, Marco, see Vizzini & al.
Cortez, Vagner G. Type studies on South American Strophariaceae: 2. Pholiota trinitensis
is transferred to Stropharia. 105: 7-10. 2008.
Costa, Ana M., see Dianese & al.
Crous, Pedro, see Braun & Crous
Cuenca, Gisela & Ricardo Herrera-Peraza. Scutellospora striata sp.nov., a newly
described glomeromycotan fungus from La Gran Sabana, Venezuela. 105: 79-87.
2008.
Cui, Bao-Kai & Yu-Cheng Dai. Wood-rotting fungi in eastern China 2. A new species
of Fomitiporia (Basidiomycota) from Wanmulin Nature Reserve, Fujian Province.
105: 343-348. 2008.
Cui, Bao-Kai, see Dai & al.
492
Cui, Bao-Kai, see Xiong & al.
Dai, Yu-Cheng, Bao-Kai Cui & Wan-Qiang Tao. Phellinus mori sp. nov. (Basidiomycota,
Hymenochaetales) from northern China. 105: 53-58. 2008.
Dai, Yu-Cheng, see Cui & Dai
Dai, Yu-Cheng, see Wei & Dai
Dai, Yu-Cheng, see Xiong & al.
Deng, Jingshi, see Su & al.
Desjardin, Dennis E., see Sulzbacher & al.
Dhingra Gurpaul S., & Aveneet P. Singh. Validation of Repetobasidiopsis and Trimitiella
(Basidiomycetes). 105: 421-422. 2008.
Di Massimo, Gabriella, see Garcia-Montero & al.
Dianese, Alexei C., Ana M. Costa & José C. Dianese. A new Pseudocercospora species
on Passiflora setaceae. 105: 1-5. 2008.
Dianese, José C., see Dianese & al.
Didukh, Marina, see Kosakyan & al.
Dissara, Yaowaluk, see Orihara & al.
Elix, John A., see Marcelli & al.
Fernandez, Roberto, see Iturriaga & al.
Franco-Molano, A.E., see Tulloss & Franco-Molano
Fulgenzi, Tara D., see Mayor & al.
Garcia-Montero, Luis G., Gabriella Di Massimo, José L. Manjon & Javier Velazquez.
Description of Tuber malenconii mycorrhizae and a comparison with mycorrhizae
of T: aestivum and T. mesentericum. 105: 407-414. 2008.
Gené, Josepa, see Castaneda Ruiz & al.
Gené, Josepa, see Silvera-Simon & al.
Ghobad-Nejhad, Masoomeh, Nils Hallenberg & Heikki Kotiranta. Additions to the
corticioids of the Caucasus from NW Iran. 105: 269-293. 2008.
Gibertoni, Tatiana Baptista, see Gomes-Silva & al.
Gleason, Mark L., see Zhai & al.
Gomes-Silva, Allyne Christina, Leif Ryvarden & Tatiana Baptista Gibertoni. Coltricia
fragilissima, a new record for Brazil. 105: 469-472. 2008.
Gonzalez, Maria C., see Rosique-Gil & al.
Goto, Bruno Tomio, Leonor Costa Maia & Fritz Oehl. Ambispora brasiliensis, a new
ornamented species in the arbuscular mycorrhiza-forming Glomeromycetes. 105:
11-18. 2008.
Guarro, Josep, see Silvera-Simon & al.
Gugliotta, Adriana de Mello, see Torres-Torres & al.
Guo, Lin, see Li & Guo
Gurgel, FE., B.D.B. Silva & I.G. Baseia. New records of Scleroderma from Northeastern
Brazil. 105: 399-405. 2008.
Guzman- Davalos, Laura, see Torres-Torres & al.
Halek, Vaclav, see Holec & Halek
493
Halici, Mehmet Gokhan. Arthonia hawksworthii sp. nov. (Ascomycota, Arthoniaceae)
on Dimelaena oreina from Turkey. 105: 89-93. 2008.
Halici, Mehmet Gokhan. Llimoniella muralicola sp. nov. (Ascomycota, Helotiaceae) on
Protoparmeliopsis muralis from western Turkey. 105: 203-206. 2008.
Hallenberg, Nils, see Ghobad-Nejhad & al.
Halling, Roy E., see Mayor & al.
Han, Yanfeng, see Liang & al.
Hao, Yue, see Su & al.
Haq, Ikram-UI, see Sultan & al.
He, Xueyou, see Liang & al.
Henkel, Terry W., see Largent & al.
Henkel, Terry W., see Mayor & al.
Heredia Abarca, Gabriela, see Castameda Ruiz & al.
Heredia Abarca, Gabriela, see Iturriaga & al.
Herrera-Peraza, Ricardo, see Cuenca & Herrera-Peraza
Holec, Jan & Vaclav Halek. Record of the rare greenhouse fungus Lepiota elaiophylla
(Agaricales, Agaricaceae) in Prague, Czech Republic, with notes on its taxonomy
and distribution. 105: 433-439. 2008.
Hughes, Karen W., see Petersen & al.
Iturriaga, Teresa, Roberto Fernandez, Rafael FE. Castafeda Ruiz, David W. Minter
& Gabriela Heredia Abarca. A new anamorphic fungus from Venezuela:
Pyriculariopsis formosa. 105: 337-342. 2008.
Iturriaga, Teresa, see Castaneda Ruiz & al.
Kasuya, Taiga, see Orihara & al.
Kendrick, Bryce, see Li & al.
Khalid, A.N., see Afshan & al.
Khalid, A.N., see Sultan & al.
Knudsen, Kerry & Jana Kocourkova. A study of lichenicolous species of Polysporina
(Acarosporaceae). 105: 149-164. 2008.
Knudsen, Kerry, see Lendemer & al.
Kocourkova, Jana, see Knudsen & Kocourkova
Kocourkova, Jana, see Lendemer & al.
Kosakyan, Anush, Marina Didukh, Yair Ur, Solomon P. Wasser & Eviatar Nevo. Lepiota
(Agaricaceae, Basidiomycota) species diversity in Israel. 105: 355-377. 2008.
Kotiranta, Heikki, see Ghobad-Nejhad & al.
Kovalenko, Alexander E., see Petersen & al.
Kuo, Michael. Morchella tomentosa, a new species from western North America, and
notes on M. rufobrunnea. 105: 441-446. 2008.
Largent, David L., M. Catherine Aime, Terry W. Henkel & Timothy J. Baroni.
The Entolomataceae of the Pakaraima Mountains of Guyana 2: Inocephalus
dragonosporus comb. nov. 105: 185-190. 2008.
494
Lendemer, James C., Jana Kocourkova & Kerry Knudsen. Studies in lichens and
lichenicolous fungi: notes on some taxa from North America. 105: 379-386.
2008. |
Li, De-Wei, Bryce Kendrick, David Spero & Claire Macdonald. Balaniopsis triangularis
sp. nov. from indoor environments. 105: 105-110. 2008.
Li, Huanyu, see Zhai & al.
Li, Zhenying & Lin Guo. Two new species and a new Chinese record of Exobasidium
(Exobasidiales) from China. 105: 331-336. 2008.
Liang, Jiangdong, see Liang & al.
Liang, Zongqi, Xueyou He, Yanfeng Han, Shouping Cai & Jiangdong Liang. A new
species of Isaria isolated from an infected locust. 105: 29-36. 2008.
Lickey, Edgar B., see Petersen & al.
Macdonald, Claire, see Li & al.
Maia, Leonor Costa, see Goto & al.
Manjon, José L., see Garcia-Montero & al.
Marcelli, Marcelo P. & Luciana S. Canéz. Novelties on Southern Brazilian Parmeliaceae.
105: 225-234. 2008.
Marcelli, Marcelo P., Michel N. Benatti e John A. Elix. New species of Parmotrema
containing protocetraric or stictic acid from the coast of Sao Paulo State,
southeastern Brazil. 105: 235-248. 2008.
Mariaca, Ramon, see Sierra & al.
Mariotti, Mauro Giorgio, see Zotti & al.
Mayor, Jordan R., Tara D. Fulgenzi, Terry W. Henkel & Roy E. Halling. Boletellus
piakaii sp. nov. and a new distribution record for Boletellus ananas var. ananas
from Guyana. 105: 387-398. 2008.
McNeill, John, see Redhead & McNeill
MeSi¢, Armin & Zdenko Tkaléec. Entoloma reinwaldii, a rare species new to Croatia.
105: 295-300. 2008.
Minnis, Andrew M. & Walter J. Sundberg. Nomenclatural note on an Asian Pluteus.
105: 165-166. 2008.
Minter, David W., see Castafieda Ruiz & al.
Minter, David W., see Iturriaga & al.
Mo, Minghe, see Su & al.
Morozova, Olga V., see Petersen & al.
Mukhtar, Hamid, see Sultan & al.
Nevo, Eviatar, see Kosakyan & al.
Niazi, A.R., see Afshan & al.
Noordeloos, Machiel & Elias Polemis. Studies in the genus Entoloma (Basidiomycetes,
Agaricales) from the Kiklades (C. Aegean, Greece). 105: 301-312. 2008.
Oehl, Fritz, see Goto & al.
Orihara, Takimichi, Taiga Kasuya, Souwalak Phongpaichit & Yaowaluk Dissara.
Radiigera tropica (Geastraceae, Geastrales), a new species from a tropical rain
forest of Thailand. 105: 111-117. 2008.
495
Pavarino, Mario, see Zotti & al.
Petersen, Ronald H., Karen W. Hughes, Edgar B. Lickey, Alexander E. Kovalenko,
Olga V. Morozova & Nadezhda V. Psurtseva. A new genus, Cruentomycena, with
Mycena viscidocruenta as the type species. 105: 119-136. 2008.
Phongpaichit, Souwalak, see Orihara & al.
Polemis, Elias, see Noordeloos & Polemis
Pratibha, J. & D.J. Bhat. New and unusual hyphomycetes from Mahabaleshwar, India.
105: 423-431. 2008.
Psurtseva, Nadezhda V. , see Petersen & al.
Putzke, Jair, see Sulzbacher & al.
Redhead, Scott A. & John McNeill. The generic name Leratiomyces (Agaricales) once
again. 105: 481-488. 2008.
Roberts, Peter. Caribbean heterobasidiomycetes: 3. British Virgin Islands. 105: 137-
147. 2008.
Rosique-Gil, Edmundo, Maria C. Gonzalez & Joaquin Cifuentes. New records of three
freshwater ascomycetes from an urban lagoon of Tabasco, Mexico. 105: 249-256.
2008.
Ruan-Soto, Felipe, see Sierra & al.
Ryvarden, Leif, see Gomes-Silva & al.
Ryvarden, Leif, see TomSovsky & Ryvarden
Saikawa, Masatoshi, see Castafieda Ruiz & al.
Shin, Hyeon-Dong, see Choi & Shin
Sierra, Sigfrido, Joaquin Cifuentes, Felipe Ruan-Soto & Ramon Mariaca. A white form
of Auricularia fuscosuccinea from Lacandonia tropical forest, Chiapas, Mexico.
105: 415-419. 2008.
Silva, B.D.B., see Gurgel & al.
Silvera-Simon, Carolina, Josepa Gené, Josep Cano & Josep Guarro. Wardomycopsis
litoralis, a new soil borne hyphomycete. 105: 195-202. 2008.
Silvera-Simon, Carolina, see Castaneda Ruiz & al.
Singh, Aveneet P., see Dhingra & Singh
Spero, David, see Li & al.
Spribille, Toby & Curtis R. Bjérk. New records and range extensions in the North
American lignicolous lichen flora. 105: 455-468. 2008.
Stadler, Marc, see Castafieda Ruiz & al.
Su, Hongyan, Yue Hao, Xiaoyan Yang, Zefen Yu, Jingshi Deng & Minghe Mo. A new
species of Dactylellina producing adhesive knobs and non-constricting rings to
capture nematodes. 105: 313-316. 2008.
Sultan, Muhammad Asim, Ikram-Ul-Hagq, A.N. Khalid & Hamid Mukhtar. Two new
anamorphic rust fungi from northern areas of Pakistan. 105: 23-27. 2008.
Sulzbacher, Marcelo A., Dennis E. Desjardin & Jair Putzke. Calathella columbiana
(Basidiomycota): new record of a cyphelloid fungus from Brazil. 105: 37-42.
2008.
Sun, Guangyu, see Zhai & al.
496
Sundberg, Walter J., see Minnis & Sundberg
Tang, Ming, see Zhai & al.
Tao, Wan-Qiang, see Dai & al.
Tkaléec, Zdenko, see Mesi¢ & Tkaléec
TomSovsky, Michal & Leif Ryvarden. Gloeoporus dichrous var. niger comb. nov. 105:
171-174. 2008.
Torres-Torres, Mabel Gisela, Laura Guzman-Davalos & Adriana de Mello Gugliotta.
Ganoderma vivianimercedianum sp. nov. and the related species, G. perzonatum.
105: 447-454. 2008.
Traverso, Mido, see Zotti & al.
Tulloss, Rodham E. & A. E. Franco-Molano. Studies in Amanita subsection Vittadiniae
1— a new species from Colombian savanna. 105: 317-323. 2008.
Tiirk, Aysen Ozdemir, see Candan & Tiirk
Ur, Yair, see Kosakyan & al.
Velazquez, Javier, see Garcia-Montero & al.
Vizzini, Alfredo, Giovanni Consiglio, Vladimir Antonin & Marco Contu. A new
species within the Gymnopus dryophilus complex (Agaricomycetes, Basidiomycota)
fom Italy. 105: 43-52. 2008.
Vizzini, Alfredo, see Zotti & al.
Wasser, Solomon P., see Kosakyan & al.
Wei, & Yu-Lian Yu-Cheng Dai. Notes on Elmerina and Protomerulius (Basidiomycota).
105: 349-354. 2008.
Xiong, Hong-Xia, Yu-Cheng Dai & Bao-Kai Cui. Perenniporia minor (Basidiomycota,
Polyporales), a new polypore from China. 105: 59-64. 2008.
Yang, Xiaoyan, see Su & al.
Yang, Zhu L., see Zhou & Yang
Yazici, Kenan & André Aptroot. Corticolous lichens of the city of Giresun with
descriptions of four species new to Turkey. 105: 95-104. 2008.
Yu, Zefen, see Su & al.
Zhai, Xiaoru, Huanyu Li, Rong Zhang, Guangyu Sun, Ming Tang, Jean C. Batzer &
Mark L. Gleason. Zygophiala (hyphomycetes) — a genus newly recorded from
China. 105: 317-322. 2008.
Zhang, Rong, see Zhai & al.
Zhou, D.Q. & Zhu L. Yang. Notes on two boletes with tiny basidioma in tropical China.
105: 473-479. 2008.
Zotti, Mirca, Alfredo Vizzini, Mido Traverso, Fabrizio Boccardo, Mario Pavarino &
Mauro Giorgio Mariotti. The macrofungal checklist of Liguria (Italy): current
survey status. 105: 167-170. 2008.
497
REVIEWERS, VOLUME ONE HUNDRED FIVE
The Editors express their appreciation to the following individuals who have, prior
to acceptance for publication, reviewed one or more of the papers appearing in this
volume.
Mehrdad Abbasi
Vagn Alstrup
Vladimir Antonin
Violeta Atienza
Timothy J. Baroni
Robert Bauer
Annarosa Bernicchia
Janusz Blaszkowski
Paola Bonfante
Uwe Braun
Peter Buchanan
Marta Cabello
Lori M. Carris
Michael A. Castellano
Marco Contu
Jerry A. Cooper
Ana Crespo
James Henry Cunnington
Yu-Cheng Dai
Cony Decock
Paul Diederich
Pradeep Kumar Divakar
Margarita Duenas
Fernando Esteve-Raventos
Javier Etayo
Roland Fox
Tatiana B. Gibertoni
Anna Guttova
Richard C. Harris
Kentaro Hosaka
Kevin D. Hyde
S.H. Iqbal
W. Bryce Kendrick
Roland Kirschner
Urmas K6ljalg
Michael Kuo
James C. Lendemer
ShiDong Li
Tai-Hui Li
XueFeng Liu
D. Jean Lodge
Eric H.C. McKenzie
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i
|
[Content continued from page 500]
| Morchella tomentosa, a new species from western North America,
|
|
and notes on M. rufobrunnea Michael Kuo 441
Ganoderma vivianimercedianum sp. nov. and the related species,
G. perzonatum Mabel Gisela Torres-Torres,
Laura Guzman-Davalos & Adriana de Mello Gugliotta 447
New records and range extensions in the North American
lignicolous lichen flora Toby Spribille & Curtis R. Bjork 455
| Coltricia fragilissima, a new record for Brazil Allyne Christina Gomes-Silva,
Leif Ryvarden & Tatiana Baptista Gibertoni 469
Notes on two boletes with tiny basidioma in tropical China
D.Q. Zhou & Zhu L. Yang 473
The generic name Leratiomyces (Agaricales) once again
Scott A. Redhead & John McNeill 481
INDICES ¢& INFORMATION
Nomenclatural novelties proposed in volume 105 489
Author index 49]
Reviewers 497
Errata 49§
500
[Content continued from inside back cover]
Studies in the genus Entoloma (Basidiomycetes, Agaricales) from the
Kiklades (C. Aegean, Greece) Machiel Noordeloos & Elias Polemis 301
A new species of Dactylellina producing adhesive knobs and
non-constricting rings to capture nematodes Hongyan Su, Yu'e Hao,
Xiaoyan Yang, Zefen Yu, Jingshi Deng & Minghe Mo 313
Studies in Amanita subsection Vittadiniae 1— a new species from
Colombian savanna Rodham E. Tulloss & A.E. Franco-Molano 317 —
Zygophiala (hyphomycetes) — a genus newly recorded from China
Xiaoru Zhai, Huanyu Li, Rong Zhang, Guangyu Sun,
Ming Tang, Jean C. Batzer & Mark L. Gleason 325
Two new species and a new Chinese record of Exobasidium
(Exobasidiales) from China Zhenying Li & Lin Guo 331.
A new anamorphic fungus from Venezuela: Pyriculariopsis formosa
Teresa Iturriaga, Roberto Fernandez, Rafael F. Castaneda Ruiz,
David W. Minter & Gabriela Heredia Abarca 337 |
Wood-rotting fungi in eastern China 2. A new species of
Fomitiporia (Basidiomycota) from Wanmulin Nature Reserve,
Fujian Province Bao-Kai Cui & Yu-Cheng Dai 343 _
Notes on Elmerina and Protomerulius (Basidiomycota)
Yu-Lian Wei & Yu-Cheng Dai 349
Lepiota (Agaricaceae, Basidiomycota) species diversity in Israel
Anush Kosakyan, Marina Didukh,
Yair Ur, Solomon P. Wasser & Eviatar Nevo 355
Studies in lichens and lichenicolous fungi: notes on some taxa
from North America James C. Lendemer,
Jana Kocourkova & Kerry Knudsen 379 |
Boletellus piakaii sp. nov. and a new distribution record for Boletellus
ananas var. ananas from Guyana Jordan R. Mayor, Tara D. Fulgenzi,
Terry W. Henkel & Roy E. Halling 387
New records of Scleroderma from Northeastern Brazil
EE. Gurgel, B.D.B. Silva & I.G. Baseia 399
Description of Tuber malenconii mycorrhizae and a comparison with
mycorrhizae of T: aestivum and T: mesentericum Luis G. Garcia-Montero,
Gabriella Di Massimo, José L. Manjon & J. Velazquez 407
A white form of Auricularia fuscosuccinea from Lacandonia tropical forest,
Chiapas, Mexico Sigfrido Sierra, Joaquin Cifuentes,
Felipe Ruan-Soto & Ram6n Mariaca 415
New and unusual hyphomycetes from Mahabaleshwar, India
J. Pratibha & D.J. Bhat 423
Record of the rare greenhouse fungus Lepiota elaiophylla
(Agaricales, Agaricaceae) in Prague, Czech Republic, with notes on
its taxonomy and distribution Jan Holec & Vaclav Halek 433 |
[Content concluded on page 499]
[Content continued from back cover]
A study of lichenicolous species of Polysporina (Acarosporaceae)
Kerry Knudsen & Jana Kocourkova 149
Nomenclatural note on an Asian Pluteus
Andrew M. Minnis & Walter J. Sundberg 165
The macrofungal checklist of Liguria (Italy): current survey status
Mirca Zotti, Alfredo Vizzini, Mido Traverso, Fabrizio Boccardo,
Mario Pavarino & Mauro Giorgio Mariotti 167
Gloeoporus dichrous var. niger comb. nov. Michal TomSovsky & Leif Ryvarden 171
Notes on Heteroconium and a new species from Venezuela
Rafael F. Castafeda Ruiz, Teresa Iturriaga, Gabriela Heredia Abarca,
David W. Minter, Josepa Gene, Marc Stadler,
Masatoshi Saikawa & Carolina Silvera-Simon 175
The Entolomataceae of the Pakaraima Mountains of Guyana 2:
Inocephalus dragonosporus comb. nov. David L. Largent,
M. Catherine Aime, Terry W. Henkel & Timothy J. Baroni 185
Reclassification of Bremia domingensis to the genus Plasmopara
as P. domingensis comb. nov. Young-Joon Choi & Hyeon-Dong Shin 191
Wardomycopsis litoralis, a new soil borne hyphomycete
Carolina Silvera-Simon, Josepa Gené, Josep Cano & Josep Guarro 195
Llimoniella muralicola sp. nov. (Ascomycota, Helotiaceae) on
Protoparmeliopsis muralis from western Turkey | Mehmet Gékhan Halici 203
Cercosporoid hyphomycetes on hosts of the Annonaceae:
Cercospora annonaceae and Isariopsis annonarum revisited
Uwe Braun & Pedro Crous 207
Novelties on Southern Brazilian Parmeliaceae
Marcelo P. Marcelli & Luciana S. Canéz 225
New species of Parmotrema containing protocetraric or stictic acid
from the coast of Sao Paulo State, southeastern Brazil
Marcelo P. Marcelli, Michel N. Benatti & John A. Elix 235
New records of three freshwater ascomycetes from an urban lagoon
of Tabasco, Mexico Edmundo Rosique-Gil,
Maria C. Gonzalez & Joaquin Cifuentes 249
New records and distribution of rust fungi from Pakistan
N.S. Afshan, A.N. Khalid & A.R. Niazi 257
Additions to the corticioids of the Caucasus from NW Iran
Masoomeh Ghobad-Nejhad, Nils Hallenberg & Heikki Kotiranta 269
Entoloma reinwaldii, a rare species new to Croatia
Armin MeSi¢ & Zdenko Tkaléec 295
[Content continued opposite on page 500]
ARE
TABLE OF CONTENTS
ue oe Dx wht :
HRs aa taerat Twila ae
A new Pseudocercospora s species on Passiflora setaceae ade
_ Alexei C. Dianese, Ana M.C a
tory
Type studies on South American vee ee pa Pholiota rinite
transferred to Stropharia Foor vig ti oe piss
mycorrhiza- PB Glomeromyezies
a. - Leonor Costa Maiz : Fr
Lichens of Malatya, Elazig, Adiyaman Piorieela of Turkey — ad sick ae
Mehmet Candan & Afien Ozdemir Ti |
Two new anamorphic r rust fungi from northern areas of Pakistan ie bie ihe yf
Muhammad Asim Sultan, Ikram-UI- -Hagq, A.N. Khalid & Hamid Mukhtar 2
A new species of Isaria isolated from an infected locust | - Zongqi Liang, APE
_ Xueyou He, Yanfeng Han, Shouping Cai & Jiangdong Liang Pebee
Calathella columbiana (Basidiomycota): new record of a cyphelloid fungus | ‘ i ‘
from Brazil Marcelo A. Sulzbacher, Dennis E. Desjardin & Jair Putzke BY
A new species within the Gymnopus dryophilus complex. (Agaricomycetes, oe gn Na
Pa ect from Italy Alfredo Vizzini, Giovanni Consiglio, —
-—- Viadimir Antonin & Marco Contu | : 4B
Phellinus mori sp. nov. (Basidiomycota, Hymenochaetales) from northern China — 3 ig
Yu-Cheng Dai, Bao- ‘Kai Cui & Wan- -Qiang Tao 5: f
Perenniporia minor (Basidiomycota, Polyporales), a new polypore from China
Hong-Xia Xiong, Yu- -Cheng Dai & Bao- Kai Cui
A taxonomic study of Heterodermia (Lecanorales, Ascomycota) in
South Korea based on phenotypic and phylogenetic analysis Xin Li Wei, iF
Heng Luo, Young Jin Koh & fie Seoun Hur ‘6
Scutellospora striata sp.nov., a newly described glomeromycotan fungus — Hi
from La Gran Sabana, Venezuela. Gisela Cuenca & Ricardo Herrera-Peraza
Arthonia hawksworthii sp. nov. (Ascomycota, Arthoniaceae) on oR
Dimelaena oreina from Turkey Mehmet Gékhan Hahei u
Corticolous lichens of the city of Giresun with descriptions of four species _ 4
new to Turkey Kenan Yazici & André Aptroot
Balaniopsis triangularis sp. Nov. rae indoor environments —-De-WeiL Li,
__ Bryce Kendrick, David Spero & Claire Macdonald d 105
Ratliigera tropica (Geastraceae, Geastrales), a new species from a tropical - ¥ ee a
rain forest of Thailand ) Takimichi Orihara, Taiga Kasuye,
Souwalak Phongpaichit & Yaowaluk Dissara ’ a i
A new genus, Cruentomycena, with Mycena viscidocruenta as the type species
Ronald H. Petersen, Karen W. rset eel B. Lick Yy
a
Caribbean he ebabaaidionehetis 3. British Virgin Linde abe
[Content continues inside back cover]
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MYCOTAXON
Volume 106, pp. 1-6 October-December 2008
First record of Discoxylaria myrmecophila
(Ascomycotina, Xylariales) from Veracruz with new reports
from Jalisco, Morelos, and Nuevo Leon (Mexico)
ROSARIO MEDEL, GASTON GUZMAN & FLORENCIA RAMIREZ-GUILLEN
rosario.medel@inecol.edu.mx gaston.guzman@inecol.edu.mx
Instituto de Ecologia, Unidad de Micologia
Apartado Postal 63, Xalapa, 91000, Veracruz, Mexico
Abstract — Fresh collections of stromata of Discoxylaria myrmecophila growing in an
ant nest in the State of Veracruz have been studied. Both teleomorph and anamorph
stages were found. A description and photographs of the “stroma” are presented. New
collections are also reported from the states of Jalisco, Morelos and Nuevo Leén.
Key words — attine ants, Hypocreodendron, distribution
Introduction
Discoxylaria J.C. Lindg. & J.E. Wright is a monotypic genus of Xylariaceae in the
Ascomycotina, with a strong relationship with ant nests of Acromyrmex Mayr
and Atta Fabricius. The teleomorph, described by Lindquist and Wright (1964)
from Argentina and Uruguay, formed in the same stroma where the anamorph
Hypocreodendron sanguineum Henn. also developed. Hennings first described
the anamorph stage from Argentina in 1897 (Rogers et al. 1995). In Mexico
this fungus was first reported from Morelos by Perez-Silva (1974) and from an
unknown locality by Guzman (1977), in both cases on nests of ants belonging
to the genus Atta. Rogers et al. (1995), who also studied both teleomorph
and anamorph stages from attine nests found in Nuevo Leon, emended both
taxa: Discoxylaria myrmecophila and Hypocreodendron sanguineum. Recently,
Guzman- Davalos et al. (2003) reported D. myrmecophila from Jalisco. Regarding
other fungi associated with Atta nests in Mexico, Romero et al. (1987) studied
Phialocladus zsoltii Kreisel from Veracruz, a fungus previously known only
from Cuba (Kreisel, 1972).
In the present paper we discuss the first record of Discoxylaria myrmecophila
and its anamorph from the State of Veracruz, based on fresh stromata found in
2 ...Medel, Guzman & Ramirez-Guillén
a nest of Atta mexicana (F. Smith). New herbarium records of D. myrmecophila
from Jalisco, Morelos, and Nuevo Leon are also presented.
Materials and methods
Hand sections of stromata mounted in KOH 5% or in Melzer’s reagent were
made to observe both the teleomorph and anamorph stages. Photographs of
fresh stromata as well as microscopic features, were taken.
Results
Discoxylaria myrmecophila J.C. Lindg. & J.E. Wright, Darwiniana 13: 139. 1964;
emend. J.D. Rogers, Y.M. Ju & E San Martin, Mycologia 87: 42. 1995. Fries. 1-10
STROMATA 20-35 mm high, erect, simple to branched. The upper part as an
enlarged disc, 5—15mm diam, convex to plane, pitted, veined oralveolate, salmon-
orange or orange-reddish. Stipe 10-25 x 2—5 mm, excluding pseudorhiza size,
with the base width up to 10 mm, surface white, soon becoming pale grayish-
rose to blackish brown from the base to the top, frequently branched, smooth
to rough because of the ostioles of the perithecia; pseudorhiza up to 30 mm
long, sometimes concrescent, attenuated toward the base, covered with black
hyphae, giving a fuzzy aspect. Context subcartilaginous, white. The mycelium
mass from the ant nests, has a sweetish smell. Perithecia embedded in the lower
part of the stipe and conidiophores developed on the disc.
Ascospores 7-8 x 3—4 um, ellipsoid-inaequilateral, hyaline to brown or
blackish, with one guttule, with an incomplete germ slit and with a hyaline
permanent perispore. Asc1 75—90 x 4-6 um, cylindric, 8-spored, uniseriate,
long stipitate, ascal pore 2 x 2 um, cuboid, blue in Melzer’s reagent. PARAPHYSES
not seen. Hyphae of the context 3-6 um wide, septate, thick-walled, dark to
light brown. ConrDIopHoRES in synemmata on the disc of the stroma. CONIDIA
8-10 (-12) x 1-2 um, holoblastic, hyaline, subclavate (slightly wider at one
extreme) or cylindric.
HABITAT: on ant-nests of Atta mexicana, in public parks or in open places of
pine-oak, several types of tropical and subtropical vegetation or in xerophytic
zones.
DISTRIBUTION: South America to North America, known from Argentina
Uruguay and Mexico.
STUDIED MATERIAL: MEXICO. JALISCO, Municipality of Guadalajara, near Barranca
de Huentitlan, Guadalajara Zoo, August 6, 1988, Guerrero 37-A (IBUG); Municipality
of Zapopan, University of Guadalajara Sport Club, July 15, 1984, Mendoza s.n. (IBUG,
XAL); Bosque La Primavera, July 18, 1985, Guzman-Davalos 2058; July 26, 1985,
Guzman-Davalos 2115 (both IBUG); University of Guadalajara, garden of Instituto
de Botanica, October 7, 1986, Guzman-Davalos 3624 (IBUG); July 1, 1997, Soto s.n.
Discoxylaria myrmecophila, new to Veracruz (Mexico) ... 3
Fics. 1-4. Discoxylaria myrmecophila. 1: Bifurcate stromata; 2: Stromata showing stipe dark color
after beginning to dry; 3: Undulating disc surface; 4: Stromata showing the white context (from
Lara 202). Scale bar = 5 mm. [Photographs: 1, 3 R. Gaitan; 2, 4 P. Callac.]
(IBUG); S of La Venta del Astillero, Rancho El Paraiso, Carrillo-Reyes 242 (IBUG);
Puente Grande, July 5, 1986, Cruz 12 (IBUG). MORELOS, Cuernavaca, July 20, 1979,
Lopez 586 (XAL); Municipality of Tepoztlan, S of Santo Domingo, October 12, 1990,
Pérez de la Rosa s.n. (IBUG). NUEVO LEON, Municipality of Guadalupe, La Pastora,
June, 1980, De la Garza s.n.; May 9, 1979, Garcia s. n. (both in XAL). VERACRUZ,
Municipality of Ixtaczoquitlan, Barranca de San Miguel, without date, Gdmez s.n.
(XAL); Municipality of Xalapa, Xalapa city, Los Lagos Park, Nov. 23, 2007, Lara 202;
Nov. 25, 2007, Lara 208 (both XAL).
Discussion
With the exception of one collection (Lopez 586, XAL), all the studied materials
lacked ascospores, although closed, empty black ostioles of perithecia were
observed on the stipe. According to Rogers et al. (1995), absence of ascospores
from the stroma appears to be common in Discoxylaria myrmecophila. Since
A Medel, Guzman & Ramirez-Guillén
Fics. 5-10. Discoxylaria myrmecophila and Hypocreodendron sanguineum. 5: Undulate surface
of the disc of the stroma; 6: Perithecial ostioles on the stipe surface; 7: Inaequilateral ascospores;
8: Asci showing ascal pore; 9: Tranverse view of H. sanguineum showing depression that forms the
undulate surface of the disc; 10: Conidia [detail] (figs. 5, 9, 10: from Lara 202; figs. 6-8: from Lopez
586. Scale bars: 5&6 = 1 mm; 7, 8 & 10 = 10 um; 9 = 25 um [Photographs: R. Medel.]
Discoxylaria myrmecophila, new to Veracruz (Mexico) ... 5
teleomorph and anamorph appear together within the same stroma, Rogers et
al. (1995) suggest that the anamorph probably appears before the teleomorph,
although the perithecia lack ascospores. Interestingly, the sweetish smell
associated with the nest’s mycelial mass in Lara 208 was first recorded by
Hennings, who studied the anamorph in culture in 1897 (Rogers et al. 1995).
Discoxylaria myrmecophila displays a disjunct geographic distribution and has
been recorded only from Argentina, Uruguay and Mexico. Surprisingly, Dennis
(1979) did not record this species, and even the fungal databases of Costa Rica
(atta.inbio.ac.cr/), Ecuador (www.mycokey.com/Ecuador) and the Caribbean
(www. biodiversity.org) do not mention this fungus. Despite more than 50 years
of field mycological work in Mexico, we have only a few collections of this fungus.
Given that this is the first record from Veracruz, where so much mycological
research has been conducted, we consider Discoxylaria myrmecophila a “rare”
fungus. However, in view of the broad distribution of ants belonging to the
Formicidae tribe Attini throughout the Americas recorded by Kempf (1972)
from the southern United States to Central America (Kempf 1972) and by
Flores-Maldonado et al. (1999) from Tamaulipas, Mexico, we expect that this
fungus has a wide distribution in subtropical America.
Acknowledgments
We express our thanks to Dr. Laura Guzman- Davalos (Universidad de Guadalajara) and
Dr. Richard Hanlin (University of Georgia) for reviewing this paper. We also thank Dr.
Jorge Valenzuela (Instituto de Ecologia) for providing information on the ants and their
geographical distribution and especially acknowledge Juan Lara Carmona (Instituto de
Ecologia) for fresh collections and Rigoberto Gaitan (Instituto de Ecologia) and Phillipe
Callac (Institute Natinale de la Recherche Agronomique, Bordeaux, France) for the
color photographs. Laura Guzman-Davalos is also thanked for the loan of Discoxylaria
myrmecophila collections from IBUG.
Literature cited
Dennis RWG. 1979. Fungus flora of Venezuela and adjacent countries, Cramer, Lehre.
Flores-Maldonado KY, Phillips SA, Sanchez-Ramos G. 1999. The myrmecophauna (Hymenoptera:
Formicidae) along an altitudinal gradient in the Sierra Madre Oriental of the Northeastern
Mexico. The Southwestern Naturalist 44: 457-461.
Guzman G. 1977. Identificacién de los hongos. Ed. Limusa, Mexico, D.F.
Guzman-Davalos L, Rodriguez O, Sanchez-Jacome MR, Chacon S. 2003. Ascomycotina conocidos
de Jalisco. Boletin IBUG (Universidad de Guadalajara) 9: 11-23.
Kempf WW. 1972. Catalogo abreviado das formigas das regiao neotropicale (Hymenop. Formicidae).
Studia Entomologica 15: 3-344.
Kreisel, H. 1972. Pilze aus Pilzgarten van Atta insularis in Kuba. Z. Allg. Mikrobiol. 12: 643-654.
Lindquist JC, Wright J. 1964. Discoxylaria género nuevo la forma perfecta de Hypocreodendron.
Darwiniana 13: 138-143.
6 ...Medel, Guzman & Ramirez-Guillén
Pérez-Silva E. 1974. Primer registro del género Discoxylaria (Pyrenomycetes) en México. Boletin de
la Sociedad Mexicana de Micologia 8: 49-52.
Rogers JD, Ju YM, San Martin FE. 1995. Discoxylaria myrmecophila and its Hypocreodendron
anamorph. Mycologia 87: 41-45.
Romero, D, Chacén $, Guzman G. 1987. Estudio y aislamiento del hongo que cultivan las hormigas
arrieras del género Atta en México. Revista Mexicana de Micologia 3: 231-248.
MYCOTAXON
Volume 106, pp. 7-13 October-December 2008
Phallus luteus comb. nov., a new taxonomic treatment
of a tropical phalloid fungus
TAIGA KASUYA
tkasuya@sakura.cc.tsukuba.ac.jp
Laboratory of Plant Parasitic Mycology,
Graduate School of Life and Environmental Sciences, University of Tsukuba
Tsukuba, Ibaraki 305-8572, Japan
Abstract — A new combination, Phallus luteus is proposed for a tropical phalloid
fungus, Dictyophora indusiata f. lutea. Macro- and microscopic characteristics are
described and illustrated based on Japanese specimens.
Key words — gasteromycetes, Japan, Phallaceae
Introduction
Phallus Junius ex L. is a gasteromycetous fungal genus of Phallaceae, Phallales
that includes morphologically variable species. Hitherto, 25 taxa of this genus
have been recognized (Calonge 2005). According to Kreisel (1996), important
morphological characteristics for taxonomy are: 1) shape and surface
configuration of the pileus; 2) presence or absence of yellow to red pigments
(carotenoids) in pileus, receptacle and indusium; and 3) presence or absence
of pinkish to purple discoloration in peridium and rhizomorphs. The presence
of an indusium has been emphasized as an important taxonomic characteristic
by several authors (Fischer 1933, Kobayasi 1938, Pilat 1958, Liu et al. 2005) so
that the genus Dictyophora Desv. has been accepted as independent in the past
based on its presence. However, Andersson (1989), Kreisel (1996), Calonge
(2005), Sarasini (2005) and Baseia et al. (2006) recognized close morphological
relationships between phalloid species with and without indusium, and they
noted that presence of an indusium is an unstable character for taxonomy.
Therefore, Dictyophora is currently put into synonymy with Phallus, following
the above authors.
Three Phallus taxa with indusia — i.e. P indusiatus Vent., P. impudicus var.
pseudoduplicatus O. Andersson, P. rubrovolvatus (M. Zang et al.) Kreisel —
have been reported from Japan (Kobayasi 1938, Ito 1959, Yoshimi & Hongo
8 ... Kasuya
1989, Kasuya et al. 2007). Moreover, another with a yellow pileus and an
indusium — Dictyophora indusiata f. lutea (= D. lutea) — has been recorded
from western Japan (Kobayasi 1938, 1942, 1965a, b; Ito 1959; Yoshimi &
Hongo 1989; Yamamoto & Yamamoto 2007), China (Liou & Hwang 1936,
Liu et al. 2005), Korea (Kobayasi 1965a) and Mexico (Guzman et al. 1990).
Kreisel (1996) and Kasuya et al. (2007) treated this fungus as “Phallus sp.” and
noted that further study was needed to clarify its taxonomic position. Below
I describe and illustrate Japanese specimens of this fungus and transfer the
species formally to Phallus. |
Materials and methods
The specimens examined in this study are deposited in the herbaria of National
Museum of Nature and Science, Tsukuba (TNS), and Miyazaki Prefectural
Museum, Miyazaki (MPM). Macroscopic characters were described by
observations on dried or fresh materials. For light microscopic observations,
free hand sections of dried or fresh specimens were mounted in water, 3%
or 5% (w/v) KOH and 30% ethanol solution on glass slides. More than forty
randomly selected basidiospores were measured under a light microscope at
1000x magnification.
Taxonomy
Phallus luteus (Liou & L. Hwang) T. Kasuya, comb. nov. Figures 1-2
MycoBAnk MB 512085
Basionym: Dictyophora lutea Liou & L. Hwang, Chinese J. Bot. 1: 89 (1936).
= Dictyophora indusiata f. lutea (Liou & L. Hwang) Kobayasi, J. Jap. Bot. 40: 179
(1965).
= Dictyophora indusiata f. aurantiaca Kobayasi, Nova Flora Japonica 2: 83 (1938).
= Phallus industiatus f. citrinus K. Das, S.K. Singh & Calonge, Bol. Soc. Micol. Madrid 31: 136
(2007).
EGG subglobose to ovoid, 30-55 x 30-60 mm, hypogeous or subhypogeous
when young, later epigeal, pale pink to reddish purple, with long, branched,
pink to reddish purple rhizomorphs up to 150 mm long. PERIDIUM three-
layered: exoperidium papery, persistent, pale pink to reddish purple, surface
smooth but slightly gyrose near the base; mesoperidium gelatinous, viscous,
thick, hyaline, pale pink to reddish purple; endoperidium membranous to
slightly fibrous, very thin, evanescent, whitish, covering upper surface of gleba. -
Fic. 1. Phallus luteus (from TNS-F-18157): A: expanding basidioma, B: mature basidioma,
C: pileus with gleba, D: indusium, E: pileus without gleba, F: two basidiomata one of them
expanding with reddish purple peridium. Bars: A, C = 20 mm; B = 45 mm; D = 5 mm; E = 10 mm;
F=18 mm.
Phallus luteus comb. nov. .
10 ... Kasuya
Fic. 2. Phallus luteus (from TNS-F-18159)
A: Pseudoparenchymatous cells of the receptacle, B: basidiospores.
Bars: A = 20 um; B = 4 um.
MATURE BASIDIOMATA 85-240 mm tall, composed of a receptacle, a pileus
covered with a gleba and an indusium. RECEPTACLE forming a pseudostipe,
70-220 mm tall, 15-25 mm diam. at apex, cylindrical, subfusoid or tapering
toward the base, spongy, hollow, surface reticulate with polygonal depressions,
white, cream to pale yellow, with pink to reddish purple peridium at the base.
PILEus conical to slightly campanulate, apex somewhat conical, truncate with
a whitish disc at the centre, 25-40 mm tall, 26-38 mm diam. at lower margin,
surface strongly reticulate, pale yellow to yellowish orange, covered with gleba;
meshes deep, polygonal to rounded, 2-5 mm diam. GLEBA mucous, olivaceous
brown to greenish black, with foetid odor. INDUstuM 60-160 mm long, papery,
fragile when dried, margin entire; dissetiment narrow, yellow to yellowish
orange; meshes polygonal to rounded, 2-7 mm diam. EXoPERIDIUM divided
into two layers; outer one up to 250-500 um thick, composed of filamentous
hyphae, 5-15 um diam., elongate, interwoven, septate, thick-walled, hyaline,
pale brown to purplish brown, with clamp-connections; inner one up to
100-250 um thick, composed of filamentous hyphae 2-4 um diam., elongate,
subparallel, gelatinous, septate, thick-walled, hyaline, with clamp-connections.
RECEPTACLE consists of pseudoparenchymatous cells 5-43 um diam., globose
to subglobose, thin-walled. Basip1A not observed. BAsip1osporEs broadly -
ellipsoid to cylindric, 3-4 x 1.5-2 um, smooth, hyaline, thin-walled, often
truncate at base.
HasitTat — On the ground or humus in evergreen woods and bamboo groves,
especially under Cryptomeria japonica, Phyllostachys pubescens, Quercus spp.
or Castanopsis spp.
Phallus luteus comb. nov. ... 11
DISTRIBUTION — Japan, China, Korea, India, tropical America (Mexico).
JAPANESE NAME — Usuki-kinugasa-take (Kobayasi 1938).
SPECIMENS EXAMINED — JAPAN. HIROSHIMA PREFECTURE, Hiroshima-shi, Asakita-
ku, Kuji, on the ground under Cryptomeria japonica woods, 23 July 2007, M. Yamate
(TNS-F-18157); YAMAGUCHI PREFECTURE, Iwakuni-shi, Misho, on the ground under
Phyllostachys pubescens grove, 14 July 2008, K. Katumoto (TNS-F-18610); Kocur
PREFECTURE, Takaoka-gun, Sakawa-cho, Mt. Kokuzo, on the ground under C. japonica
woods, 4 July 2007, Y. Yamamoto (TNS-F-18158); Kami-shi, Kahoku-cho, Ioroi, on the
ground under C. japonica woods, 7 July 2007, R. Yamamoto (TNS-F-18159); MryvAzakI
PREFECTURE, Higashi-usuki-gun, Morotsuka-son, 21 September 1976, Y. Doi (TNS-
F-227340); Higashi-usuki-gun, Saigo-son, Ohara, on the ground under Quercus spp.,
10 October 1998, S. Kurogi, B-E-27 (MPM); Nishi-morogata-gun, Takahara-cho, Oike,
on the ground under evergreen forest, July 1999, S. Kurogi, B-E-2449 (MPM); Higashi-
morogata-gun, Kunitomi-cho, Kusumi, on the ground under evergreen forest, 11
October 1999, S. Kurogi, B-E-4473 (MPM); Kobayashi-shi, Kiuragi, 12 October 1999, S.
Kurogi, B-E-2446 (MPM); Higashi-morogata-gun, Aya-cho, Kawanaka, on the ground
under evergreen forest, 19 September 2000, S. Kurogi, B-E-509 (MPM); Nobeoka-shi,
Mt. Mukabaki, on the ground under evergreen forest, July 12 2003, S. Kurogi, B-E-2129
(MPM); Miyazaki-shi, Mt. Boroishi, on the ground under evergreen forest, 10 September
2006, S. Morimoto, B-E-4099 (MPM).
Discussion
Morphological characters of the specimens examined agree well with the
previous descriptions of the present fungus (Liou & Hwang 1936, Kobayasi
1938, 1942, 1965a, b, Ito 1959, Yoshimi & Hongo 1989, Guzman et al. 1990,
Liu et al. 2005, Yamamoto & Yamamoto 2007). This fungus was defined by a
yellowish reticulate pileus, yellow indusium, and pale pink to reddish purple
peridium with rhizomorphs. These morphological characteristics suggest
its placement in Phallus subgen. Phallus sect. Flavophallus Kreisel (Kreisel
1996). It clearly differs from P. indusiatus (= D. indusiata) in its yellow pileus
and indusium, and pale pink to reddish purple peridium with rhizomorphs.
Therefore, D. indusiata f. lutea is accepted as an independent species, P. luteus.
The recently described Indian fungus, P indusiatus f. citrinus (Das et al.
2007), is morphologically very similar to P luteus in having yellow pileus and
indusium, vinaceous to dark vinaceous peridium. Original description and
color figures of P. indusiatus f. citrinus (Das et al. 2007) represent well the
present fungus in both macroscopic and microscopic features. Therefore, I
conclude they are identical. Phallus luteus having priority over P. indusiatus f.
citrinus, the name should be adopted as the correct one for the present fungus
(Art. 11.4, ICBN Vienna Code).
There are seven other known Phallus species in section Flavophallus (Kreisel
1996). Among them, P. callichrous (Moller) Lloyd, P multicolor (Berk. &
Broome) Cooke, and P. cinnabarinus (W.S. Lee) Kreisel are morphologically
12 ... Kasuya
similar to P. luteus in possession of indusium and purplish eggs. However, its
whitish indusium clearly distinguishes P. callichrous (Kobayasi 1942). Phallus
multicolor is also distinguishable from P luteus by its orange, pink to red pileus
and indusium and slightly longer (3.5-4.5 x 1.5-2 um; Cunningham 1944)
basidiospores. Further, P. cinnabarinus is clearly different from P. luteus because
the former has a cinnabar-red to brilliant red pileus and indusium (Lee 1957).
The four other species in sect. Flavophallus — P. flavocostatus Kreisel, P. tenuis
(E. Fisch.) Kuntze, P. sulphureus Lohwag and P. formosanus Kobayasi — differ
from P luteus in their white to yellowish peridium, yellow to pink receptacles
and the absence of an indusium (Kobayasi 1938, Li et al. 2004, 2005).
Phallus tenuissimus T. H. Li et al., recently described from China (Li et al.
2005), also has a yellow, reticulate pileus and a pale yellow receptacle with a
white to yellowish peridium. Although P. tenuissimus is probably placed in
section Flavophallus, it is clearly distinguished from P luteus by its peridium
colour and the absence of an indusium.
Acknowledgments
I express my sincere thanks to Prof. Francisco D. Calonge of Real Jardin Botanico,
Madrid, to Dr. Zhu L. Yang of Kunming Institute of Botany of the Chinese Academy of
Sciences, and to Dr. Gast6n Guzman of Instituto de Ecologia, Mexico, for their valuable
suggestions for preparing this paper. I am also grateful to Dr. Tsuyoshi Hosoya of
National Museum of Nature and Science, to Dr. Ken Katumoto of Yamaguchi Prefecture,
and to Mr. Shuichi Kurogi of Miyazaki Prefectural Government, for the accession to
the important specimens. For collecting specimens, I thank Dr. Yukinori Yamamoto of
Kochi Prefecture and Ms. Machiko Yamate of Hiroshima Prefecture.
References
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219-241.
Baseia IG, Maia LC, Calonge FD. 2006. Notes on Phallales in the Neotropics. Bol Soc Micol Madrid
30: 87-93. |
Calonge FD. 2005. A tentative key to identify the species of Phallus. Bol Soc Micol Madrid 29:
9-18;
Cunningham GH. 1944. The Gasteromycetes of Australia and New Zealand. J. McIndoe,
Dunedin.
Das K, Singh SK, Calonge FD. 2007. Gasteromycetes of Western Ghats, India: I. A new form of
Phallus indusiatus. Bol Soc Micol Madrid 31: 135-138.
Fischer E. 1933. Reihe Gasteromyceteae. In: Die Natiirlichen Pflanzenfamilien Band 7a (eds. A
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Ito S. 1959. Mycological flora of Japan. Vol.2, No. 5. Yokendo, Tokyo.
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Kreisel H. 1996. A preliminary survey of the genus Phallus sensu lato. Czech Mycol 48: 273-281.
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Li TH, Song B, Liu B. 2004. Three taxa of Phallaceae in HMAS, China. Fungal Diversity 11: 123-
IE
Li TH, Liu B, Song B, Deng WQ, Zhou TX. 2005. A new species of Phallus from China and
P. formosanus, new to the mainland. Mycotaxon 91: 309-314.
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Liu B, Fan L, Li J, Li T, Song B, Liu J. 2005. Sclerodermatales, Tulostomatales, Phallales et Podaxales.
Flora fungorum Sinicorum Vol. 23. Science Press, Beijing.
Pilat A. 1958. Phallaceae. In: Flora CSR B-1, Gasteromycetes (ed. A Pilat). Ceskoslovenské
Akademie, Praha: 50-79.
Sarasini M. 2005. Gasteromiceti epigei. A. M. B. Fondazione Centro Studi Micologici, Vicenza.
Yamamoto Y, Yamamoto R. 2007. Notes on gasteroid fungi of Shikoku (1). Bot Kochi Pref 20:
91-125.
Yoshimi S, Hongo T. 1989. Gasteromycetidae. In: Colored illustrations of mushrooms of Japan Vol.
II (eds. R Imazeki, T Hongo). Hoikusha, Osaka: 193-228.
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MY COTA XON
Volume 106, pp. 15-27 October-December 2008
Conidial fungi from semi-arid Caatinga biome of Brazil.
New and interesting Dictyochaeta species
ALISSON CARDOSO RODRIGUES DA CRUZ
alissonbio@yahoo.com.br
SHEILA MIRANDA LEAO-FERREIRA
Sheilalleao@yahoo.com.br
FLAVIA RODRIGUES BARBOSA
faurb10@yahoo.com.br
&
Luis FERNANDO PASCHOLATI GUSMAO
lgusmao@uefs.br
Departamento de Ciéncias Biologicas, Laboratorio de Micologia
Universidade Estadual de Feira de Santana
BR116 KM03, 44031-460, Feira de Santana, BA, Brazil.
Abstract - During preparation of an inventory of dematiaceous hyphomycetes from
dead plant material in six areas of extreme biological importance of Caatinga biome,
some species of Dictyochaeta were found. Dictyochaeta caatingae and D. intermedia are
proposed as new species. Dictyochaeta britannica, D. microcylindrospora, D. obesispora,
and D. setosa are new records to the American continent, while D. heteroderae is new to
the neotropics and D. triseptata is new to Brazil. These species are described, discussed
and illustrated. A key to known species collected in Brazil is provided.
Key words - litter fungi, systematic
Introduction
The genus Dictyochaeta was proposed by Spegazzini (1923), with D. fuegiana
as the type species. An historical review of related genera, such as Codinaea
Maire and Dictyochaetopsis Aramb. & Cabello were made by Kuthubutheen
& Nawawi (1991a) and Whitton et al. (2000). More recently, two additional
species have been described, D. multifimbriata R. Kirschner & Chee J. Chen,
with reniform, 0-septate conidia, and D. curvispora L. Cai, et al., with curved,
asetulate conidia (Kirschner & Chen 2002, Cai et al. 2004).
16 ... Cruz & al.
Six species of Dictyochaeta occurring on decaying plant material in Brazil
have hitherto been collected: D. eucalypti (B. Sutton & Hodges) Whitton et al.,
D. septata (B. Sutton & Hodges) Whitton et al., D. simplex (as Codinaea, Sutton
& Hodges 1975), D. fertilis (Grandi 1990), D. novae-guineensis (Grandi 1998)
and D. pluriguttulata (Barbosa et al. 2007).
Materials and methods
The semi-arid Caatinga biome was described in previous papers (Cruz et al.
2007a,b, Marques et al. 2007). Samples of plant litter were placed in separate
paper bags and taken to the laboratory. The litter was then incubated at 25° C in
Petri dishes, within moist chambers, consisting of plastic boxes (50 L capacity)
containing 200 ml of sterile water plus 2 ml of glycerol (Castafieda-Ruiz 2005).
The samples were examined at regular intervals for the presence of microfungi.
Mounts were prepared in polyviny! alcohol-glycerol (8.0 g in 100 ml of water,
plus 5 ml of glycerol) and measurements made at a magnification of x1000.
Taxonomic part
Dictyochaeta caatingae A.C. Cruz & Gusmao anam. sp. nov. Fics 1-2
MycoBank MB511940
COLONIAE effusae, brunneae. MYCELIUM plerumque in substrato immersum. HYPHAE
septate, ramosae, laeviae, brunneae. SETAE solitariae vel 2-3 fasciculatae, erectae, rectae
vel flexuosae, 9-15-septatae, laeves, subulatae, basim versus atrobrunneae, apicem versus
brunneae vel pallide brunneae, 160-295 x 4.5—6 ym, ad basim dilatata 7.5—9.5 ym latae.
CONIDIOPHORA Solitaria vel 2-5 fasciculata, macronematosa, mononematosa, erecta,
recta vel flexuosa, 1-4-septata, simplicia, laevia, basim versus pallide brunnea, apicem
versus subhyalina, 37-89 x 3.5—6 um. CELLULAE CONIDIOGENAE monophialidicae, raro
polyphialidicae in conidiophoris incorporatae, determinatae, raro cum proliferationibus
sympodialis, cylindricae, pallide brunneae vel hyalinae, 12-48 x 3.5—6 um. CONIDIA 3-
septata, plus minusve falcata, laevia, hyalina, in massam mucosum, 27-35.5 x 3.3-5.5
um; plerumque utrinque setulis simplicibus, 0.5—2.5 um longis praedita. TELEOMORPHOSIS
ignota.
Hotortypus: HUEFS 129377. BRAZIL. BAuta: Senhor do Bonfim, in foliis angiospermae
ignotae, 10.X.2006, coll. A.C.R. Cruz.
ETYMOLOGY: caatingae, in reference to the Caatinga biome.
CoLony effuse, brown. MycELIuM immersed in substrate. HYPHAE septate,
branched, smooth, brown. SETAE solitary or in groups of 2-3, erect, straight
or flexuous, 9-15-septate, smooth, subulate, brown to pale brown, 160-295.
x 4.5—6 um; basal cell swollen to 7.5—9.5 um. CONIDIOPHORES solitary or in
groups of 2-5, macronematous, mononematous, straight or flexuous, 1-4-
septate, simple, smooth, brown to subhyaline at apex, 37-89 x 3.5-6 um.
CONIDIOGENOUS CELLS monophialidic, determinate or rarely polyphialidic,
then sympodial, integrated, cylindrical, pale brown to hyaline, 12—48 x 3.5-6
Dictyochaeta spp. from Caatinga biome (Brazil) ... 17
tum; collarette, 1-3 x 1-2 um. ConrpIA 3-septate, falcate to allantoid, smooth,
hyaline, in slimy mass, 27—35.5 x 3.3-5.5 um. SETULAE filiform 1 or 2, differing
in number at each end, 0.5—2.5 um long. TELEOMORPH unknown.
CoMMENTs: Dictyochaeta matsushimae (Hewings & J.L. Crane) Whitton et
al., is the only species that has 3-septate conidia and sterile setae. However,
the conidia of D. matsushimae are shorter, although the setulae are longer,
than those of D. caatingae. D. macrospora Kuthub. & Nawawi (Kuthubutheen
& Nawawi 1991b) and D. triseptata (Castaheda-Ruiz 1986) are similar to the
proposed new species, but possess consistently polyphialidic conidiophores
and longer conidia and setulae (Table 1).
TaBLE 1. Comparison of Dictyochaeta species morphologically similar to D. caatingae
CONIDIA
SPECIES SETAE CONIDIOGENESIS
SEPTA (#) SIZE (um) SETULAE (um)
D. brevisetula Present polyphialidic 1-3 21-24.5 x 3-3.7 1.4-2.7
D. caatingae Present monopiia eich 3 27-35.5 x 3.3-5.5 0.5-2.5
polyphialidic
monophialidic/ 0 10-18 x 1-2 absent
Becrmorpaa ea polyphialidic 3 22-28 x 7-8 5-6
D. macrospora Absent polyphialidic 3 38-47 x 5-7 22-30
D. matsushimae __ Present polyphialidic (1-2)-3 20-27 x 3.3-5.4 5.5-7.8
D. septata Present polyphialidic 1-3 (14.5-)17-23 x 2 5.5-10
D. triseptata Absent polyphialidic 3 21-30 x 6-7.5 3-6
D. variabilis Absent monophialidic 1-3 30-40 x 4.5-5.7 17-23
Dictyochaeta intermedia Gusmao & S.M. Leao, anam. sp. nov. FIGS 3-4
MycoBANK MB511941
COLONIAE effusae, brunneae. MYCELIUM plerumque in substrato immersum. HYPHAE
septate, ramosae, laeviae, brunneae. SETAE solitariae, erectae, rectae, 9-15-septatae,
laeves, subulatae, basin versus atrobrunneae, apicem versus brunneae vel pallide brunneae,
150-354 x 6-9 um; cellula penultima fusca, 25—47.5 um longa. CONIDIOPHORA solitaria
vel 2-5 fasciculata, macronematosa, mononematosa, erecta, recta vel flexuosa, 2-7-
septata, simplicia, laevia, basim versus pallide brunnea, apicem versus subhyalina, 30-90
x 3-6 um. CELLULAE CONIDIOGENAE monophialidicae, in conidiophoris incorporatae,
determinatae, cylindricae, cum collarulo distincte, 1.8-2.4 um longa, pallide brunneae
versus hyalinae. Conip1A 0-septata, falcata, laevia, pluriguttulata, hyalina, in massam
mucosum, 18-25 x 1.5-4 um; plerumque utrinque setulis simplicibus, 3.5-6.5 um longis
praedita. TELEOMORPHOSIS ignota.
Hototypus: HUEFS 114791. BRAZIL. BAuta: Senhor do Bonfim, in foliis angiospermae
ignotae, 10.X.2006, coll. S.M. Ledo-Ferreira.
ETYMOLOGy: intermedia, in reference to position between two previous described
species.
18 ... Cruz & al.
25um
Fics 1-2. Dictyochaeta caatingae. 1. Setae and conidiophore. 2. Conidia.
Fics 3-4. Dictyochaeta intermedia. 3. Seta and conidiophores. 4. Conidia.
CoLony effuse, brown. MycELIUM immersed in substrate. HYPHAE septate,
branched, smooth, brown. SETAE solitary, erect, straight, 9-15-septate, smooth,
brown at the base, brown to pale brown at apex, 150-354 x 6—9 um; penultimate ~
cell dark brown, 25-47.5 um long. CONIDIOPHORES arising solitary or in
groups of 2-5, associated with base of setae, macronematous, mononematous,
erect, straight or flexuose, 2—7-septate, simple, smooth, pale brown at the bases,
subhyaline at apex, 30-90 x 3-6 um. CONIDIOGENOUS CELLS monophialidic,
determinate, integrated, cylindrical, conspicuous collarettes 1.8-2.4 um long,
pale brown to hyaline. Conrp1A 0-septate, falcate, simple, smooth, pluriguttulate,
Dictyochaeta spp. from Caatinga biome (Brazil) ... 19
in slimy mass, hyaline, 18-25 x 1.5-4 um. SETULAE simple at each end, 3.5-6.5
um long. TELEOMORPH unknown.
CoMMENTSs: The combination of characters including setae with one cell darker
and conidia pluriguttulate are found only in three species of Dictyochaeta
hitherto described: Dictyochaeta anamorph of Chaetosphaeria pulchriseta
S. Hughes et al. (Hughes & Kendrick 1968), D. pluriguttulata and D. vittata
Kuthub. & Nawawi (Kuthubutheen & Nawawi 1991b). The inconspicuous
setulae and asymmetrical morphology of conidia in D. pluriguttulata are
sufficient to separate it from D. intermedia. The new species is most similar
to Dictyochaeta anamorph of Chaetosphaeria pulchriseta and D. vittata in
the general morphology but the size of conidia and setulae are intermediate
between the two known species (Table 2).
TABLE 2. Comparison of Dictyochaeta species morphologically similar to D. intermedia
SETAE (um) ConipiA (um)
SPECIES
SIZE DARK CELL SIZE SETULAE
D. intermedia 150-354 x 6-9 25-47.5 18-21 x 1.5-2.5 3.5-6.5
D. pluriguttulata up to 220 x 5-7 20-50 20-27% 2-25), inconspicuous
D. anamorph of
Chaetosphaeria up to 360 x 5.6-7 36-81 23-29 x 2.4-3 up to 1.8
pulchriseta
D. vittata up to 350 x 6-8 20-35 25-32 xX 2.5-3 10-12
Dictyochaeta britannica (M.B. Ellis) Whitton, McKenzie & K.D. Hyde,
Fungal Diversity 4: 137. 2000. FIGS 5-6
Bas.: Codinaea britannica M.B. Ellis, More Dematiaceous Hyphomycetes: 472. 1976.
Syn.: Menispora britannica (M.B. Ellis) P.M. Kirk, Mycotaxon 23: 334. 1985.
SETAE solitary, erect, straight or flexuous, septate, subulate, smooth, brown
to subhyaline, 60-345 x 3—6 um; apex acerose to rounded. CONIDIOPHORES
macronematous, mononematous, solitary or in groups of 2-5 arising from the
base of setae, straight or flexuous, septate, simple, smooth, brown to pale brown,
49.5-79.5 x 3-4.5 um. CONIDIOGENOUS CELLS monophialidic or polyphialidic,
integrated, sympodially proliferating, conspicuous collarettes, pale brown.
ConrpiA solitary, 0-septate, lunate, simple, smooth, in slimy mass, hyaline,
12.5-19 x 1—2.5 um. SETULAE filiform on each end, 7—10.5 um.
SPECIMENS EXAMINED: BRAZIL. BAHIA: Jeremoabo, “Mata das Pororocas, on
decaying bark, 17/1/2007, A.C.R. Cruz, HUEFS 129375; Paulo Afonso, on dead leaves
of unidentified dicotyledon, 31/1/2007, A.C.R. Cruz, HUEFS 129376.
KNOWN DISTRIBUTION: United Kingdown, Kenya (Ellis 1976, Kirk 1985).
Comments: Dictyochaeta britannica is similar to D. australiensis (B. Sutton)
Whitton et al., but has wider conidia and possesses a radially lobed cell at the
base of the setae. The dimensions of conidia are consistent with the descriptions
20 ... Cruz & al.
given by Ellis (1976) and Kirk (1985); however, the setulae of the examined
specimens are longer than the material studied by Kirk (1985). D. britannica is
recorded for the first time for the American continent.
Dictyochaeta heteroderae (Morgan-Jones) Carris & Glawe,
Mycotaxon 33: 23. 1988. Fics 7-8
Bas.: Codinaea heteroderae Morgan-Jones, Mycotaxon 14(1): 175. 1982.
SETAE absent. CONIDIOPHORES macronematous, mononematous, solitary,
straight or slightly flexuous, septate, simple, smooth, pale brown to brown, paler
towards the apex, 75-195 x 3—3.5 um. CONIDIOGENOUS CELLS monophialidic
or polyphialidic, integrated, terminal or intercalary, percurrently proliferating,
conspicuous collarettes, pale brown, 20—27.5 x 2.0-2.5 um. Conrp1a solitary,
0-septate, ellipsoid to allantoid, gently curved, simple, smooth, in slimy mass,
hyaline, 6—9.5 x 1—2.4 um. SETULAE absent.
SPECIMEN EXAMINED: BRAZIL. BAHIA: Santa Terezinha, Serra da Jibdia, on decaying
twig, 10/1/2008, ER. Barbosa, HUEFS 122284.
KNOWN DISTRIBUTION: Australia (Matsushima 1989), United States of America
(Morgan-Jones 1982).
CoMMENTs: Dictyochaeta heteroderae resembles D. coffeae (Maggi & Persiani)
Whitton et al. by the percurrent proliferation of the conidiogenous cells but
differs mainly in the absence of setulae at the end of conidia (Maggi & Persiani
1984). The specimen examined has conidia smaller than the material studied
by Morgan-Jones (1982) (9-13 x 3-4 um) and Matsushima (1989) (10-16 x
3-5 um). The species is recorded for the first time for the neotropics.
Dictyochaeta microcylindrospora Whitton, McKenzie & K.D. Hyde,
Fungal Diversity 4: 141. 2000. Figs 9-11
SETOSE CONIDIOPHORES, solitary, erect, straight, septate, smooth, brown at
the base, paler at apex, 93-123 x 2.5-3 um. CONIDIOPHORES macronematous,
mononematous, arranged in groups of 1-4 arising from the base of setae, straight,
septate, simple, smooth, pale brown, 32—37.5 x 1.5—2.5 um. CONIDIOGNEOUS
CELLS monophialidic, integrated, terminal, determinate, pale brown; collarettes
conspicuous, 2.4—3 um long. Conrp1A solitary, 0-septate, cylindrical, simple,
smooth, in slimy mass, hyaline, 4.2—7.8 x 1—1.2 um; SETULAE absent.
SPECIMEN EXAMINED: BRAZIL. BAHIA: Santa Terezinha, Serra da Jibdia, on dead leaves
of Clusia melchiorii Gleason, 09/II/2006, ER. Barbosa, HUEFS 123308.
KNOWN DISTRIBUTION: China (Whitton et al. 2000).
COMMENTS: Dictyochaeta microcylindrospora is closely to D. cylindrospora
(Morgan-Jones & E.G. Ingram) Whitton et al. but differs by the smaller conidia.
The species was recently found on dead leaves of Pandanus furcatus Roxb.
Dictyochaeta spp. from Caatinga biome (Brazil) ... 21
a A
2S \\ = bi i
{| al rf
< - \ ST
rs Gh} lee
8G
Figs 5-6. Dictyochaeta britannica. 5. Seta and conidiophores. 6. Conidia.
Figs 7-8. Dictyochaeta heteroderae. 7. Conidiophore. 8. Conidia.
Fics 9-11. D. microcylindrospora. 9. Setose conidiophore and conidiophores.
10. Detail of conidiogenous cell. 11. Conidia.
(Whitton et al. 2000). The species is recorded for the first time for the American
continent.
22"... Cruz al:
Dictyochaeta obesispora (S. Hughes & W.B. Kendr.) Whitton, McKenzie &
K.D. Hyde, Fungal Diversity 4: 145. 2000. FIGS 12-13
Bas.: Codinaea obesispora S. Hughes & W.B. Kendr., N.Z. Jl Bot. 6: 351. 1968.
SYNNEMATA Straight or flexuous, septate, simple, dark brown, pale brown at apex,
300-980 x 10.5-15 um. ConrDIOPHORES densely grouped, septate, branched
at apex, smooth, brown. CONIDIOGENOUS CELLS, monophialidic, terminal,
integrated, determinate, pale brown to subhyaline, collarettes inconspicuous.
Conipi solitary, 0-septate, lunate, smooth, in slimy mass, hyaline, 12—15.5 x
3.5—5.5 um. SETULAE filiform at each end, 1.2—1.8 um.
SPECIMEN EXAMINED: BRAZIL. BAHIA: Senhor do Bonfim, “Serra da Maravilha’, on
decaying bark, 11/X/2006, A.C.R. Cruz, HUEFS 129373.
KNOWN DISTRIBUTION: New Zealand (Hughes & Kendrick 1968).
COMMENTS: Only two species of Dictyochaeta have synnematal conidiomata. D.
obesispora has synnemata formed by up to 45 conidiophores and D. dendroidea
Kuthub. by up to 25 conidiophores. The conidia of D. dendroidea are smaller
than D. obesispora (Kuthubutheen 1987). Conidia and setulae for the examined
material are smaller than recorded for the type material (17-21 x 5.4-6.2 um
and 6.4 um long, respectively). Hughes & Kendrick (1968) referred to the
presence of polyblastic conidiogenous cells; however, in the Brazilian material
only monoblastic conidiogenous cells were observed. This discrepancy may be
related to differences in the age of material. This species is recorded for the first
time for the American continent.
Dictyochaeta setosa (S. Hughes & W.B. Kendr.) Whitton, McKenzie &
K.D. Hyde, Fungal Diversity 4: 148. 2000. FIGS 14-15
Bas.: Codinaea setosa S. Hughes & W.B. Kendr., N.Z.Jl. Bot. 6: 360. 1968.
SETOSE CONIDIOPHORES, solitary, erect, straight, septate, smooth, brown at base,
pale brown at apex, 246-360 x 4.5-6 um. CONIDIOPHORES macronematous,
mononematous, arising solitary or associated with base of setae, straight,
septate, simple, smooth, pale brown, 84-189 x 3-4.5 um. CONIDIOGENOUS
CELLS monophialidic, integrated, terminal, determinate, pale brown. CONIDIA
solitary, 1-septate, falcate, simple, smooth, in slimy mass, hyaline, 22.5-31.5 x
1.5-2.2 um. SETULAE absent.
SPECIMEN EXAMINED: BRAZIL. BAHIA: Santa Terezinha, Serra da Jibdia, on dead leaves
of Clusia melchiorii, 26/1V/2006, ER. Barbosa, HUEFS 1233555.
KNOWN DISTRIBUTION: England (CABI Bioscience Databases), India (Patil et al. 1991),
New Zealand (Hughes & Kendrick 1968).
COMMENTS: Dictyochaeta setosa is similar to D. ixorae (M.S. Patil et al.) Whitton
et al., but has larger conidia (Patil et al. 1991). Hughes & Kendrick (1968)
described polyblastic conidiogenous cells; however, in examined material only
monoblastic cells was observed. The dimensions of setose conidiophores and
Dictyochaeta spp. from Caatinga biome (Brazil) ... 20
wings
Fics 12-13. Dictyochaeta obesispora. 12. Synnemata. 13. Conidia.
Fics 14-15. Dictyochaeta setosa. 14. Setose conidiophore and conidiophore. 15. Conidia.
Fics 16-17. Dictyochaeta triseptata. 16. Conidiophores aggregated with different
development stages. 17. Conidia.
24 ... Cruz & al.
conidiophores are bigger than the original description (Hughes & Kendrick
1968). The species is recorded for the first time for the American continent.
Dictyochaeta triseptata (Matsush.) R.F. Castafieda,
Fungi Cubenses IJ, 8: 1986. FIGS 16-17
Bas.: Codinaea triseptata Matsush., Matsush. Mycol. Mem. 2: 4. 1981.
SETAE absent. CONIDIOPHORES macronematous, mononematous, aggregated,
rare solitary, with different development stages, straight or flexuous, smooth,
dark brown to pale brown at apex, 175-272 x 4.5-9 um. CONIDIOGENOUS
CELLS polyphialidic, integrated, sympodial and percurrent proliferating, pale
brown, conspicuous collarettes. CONIDIA solitary, 3-septate, elliptic to lunate,
simple, smooth, in slimy mass, hyaline, 18—25.5 x 4.5—7 um. SETULAE filiform
at each end, 6-9 um.
SPECIMEN EXAMINED: BRAZIL. BAHIA: Jeremoabo, “Mata das Pororocas’, on dead
leaves of unidentified dicotyledon, 18/1/2007, A.C.R. Cruz, HUEFS 129374.
KNOWN DISTRIBUTION: Cuba (Castafieda-Ruiz 1986), Micronesia (Matsushima 1981),
Guyana (Cannon & Simmons 2002) and Malaysia (CABI Bioscience Databases).
CoMMENTs: Dictyochaeta triseptata is morphologically similar to D. dimorpha
(Toyaz. & Udagawa) Whitton et al, and D. macrospora. D. triseptata is
differentiated from D. dimorpha, which has dimorphic conidiophores
and conidia, and from D. macrospora, which has larger conidia and setulae
(Kuthubutheen & Nawawi 1991b). The material examined has smaller conidia
than observed by Castafieda-Ruiz (1986); however, it is comparable with
observations by Matsushima (1981). Another interesting observation is the
conidiophores with different development stages (Fic. 16), information not
noted previously for this species.
Other species found in semi-arid regions
Dictyochaeta fertilis (S. Hughes & W.B. Kendr.) Hol.-Jech.,
Folia geobot. phytotax. 19: 426. 1984.
SPECIMENS EXAMINED: BRAZIL. Banta: Senhor do Bonfim, on decaying fruit, 28/
TX/2006, A.C.R. Cruz, HUEFS 129370; Senhor do Bonfim, on decaying fruit, 4/X/2006,
A.C.R. Cruz, HUEFS 129371.
Dictyochaeta novae-guineensis (Matsush.) A.I. Romero,
Boln Soc. Argent. Bot. 22: 76. 1983.
SPECIMENS EXAMINED: BRASIL. BAHIA: Campo Formoso, on decaying bark, 4/X/2006,
A.C.R. Cruz, HUEFS 129365; Senhor do Bonfim, on dead leaves of unidentified
dicotyledon, 5/X/2006, A.C.R. Cruz, HUEFS 129366; Senhor do Bonfim, on dead leaves
of unidentified dicotyledon, 18/X/2006, A.C.R. Cruz, HUEFS 129367; PERNAMBUCO:
Buique, on dead leaves of unidentified dicotyledon, 31/VIII/2006, A.C.R. Cruz, HUEFS
129368; Buique, on dead leaves of unidentified dicotyledon, 8/IX/2006, A.C.R. Cruz,
HUEEBS 129369.
Dictyochaeta spp. from Caatinga biome (Brazil) ... 25
Dictyochaeta pluriguttulata Kuthub. & Nawawi, Mycol. Res. 95: 1212. 1991.
SPECIMEN EXAMINED: BRAZIL. PERNAMBUCO: Buique, on dead leaves of unidentified
dicotyledon, 18/VIII/2006, A.C.R. Cruz, HUEFS 129372.
Dictyochaeta simplex (S. Hughes & W.B. Kendr.) Hol.-Jech.,
Folia geobot. phytotax. 19: 434. 1984.
SPECIMENS EXAMINED: BRASIL. Banta: Paulo Afonso, on decaying fruit, 28/1/2007,
A.C.R. Cruz, HUEFS 129362; PERNAMBUCO: Buique, on dead leaves of unidentified
dicotyledon, 28/VIII/2006, A.C.R. Cruz, HUEFS 129363; Buique, on dead leaves of
unidentified dicotyledon, 8/1X/2006, A.C.R. Cruz, HUEFS 129364.
Key to Dictyochaeta species collected in Brazil
mame OTICIODNOLES SYVNNeMalOUs eel tre rene, eae ane eae eis. ake teat D. obesispora
POR CODIGIOD NOTES ODONEMatOls meme sere eet ie IN toes. .eeee e e ee 2
Paa@Onidiaewithoutisctt jceeummne fenr e ue arnt t eats Aura dae ae ha 3
mba Onidia with sete lac sea mmre ee sees ans eg en: 3 Al ye noe ER Le 5
baw @onidiadion.septate™ 2o.saeenr ten eee. ete Ld tat ake ee, een a I, 4
Slmeonidia.t-septates2 2.9311 axa Do LUN ae Ae ee Meee tae ae ee D. setosa
pam oldia cylindrical, 4.27.8 x41 12 ieee te ee eee, D. microcylindrospora
4b. Conidia ellipsoid to allantoid, 6-9.5 x 1-2.4um.................. D. heteroderae
BAmUPOnIGI NOM =SCDLALC tea" Merrett ree cent ee ee Lee ee Ga a i ec te ee. 6
BPO OliCiase Dial Cr mer cl eer wet OTe eae ANNE, ewe uta ae heee wc 10
Gamoc acroLseLose CONIGIOPNOre DIeseNts ame me ate ee thee Meee es pene en ete 7
Bisa ClAC AD SCI) beeen x ee Bice ne ae eas ie ee ee IN hte s NOR wad ate D. simplex
PAM SCLOSE'C ONICIOD NOL ae ei aei eae mere eae oe eens creo ee ore eee ee D. fertilis
DaSelic mee et se eee nO) My: Sanwa) KU ee ae ED Oe 8
Sam@onidia pluiicuttl ate were esc ee ce eee ee ee ee ree er 9
Sba@onidianon-outtilatee| 225-19 tale Dec eee seis gee eer D. britannica
9a. Conidia asymmetrical, 20-27 x 2-2.7 um setulae inconspicuous. . D. pluriguttulata
9b. Conidia symmetrical, setulae conspicuous, 3.5-6.5 um long......... D. intermedia
LOamOnidiay =septaten we. ye. ok | ea ae dee casts one nie ee ae eh erent Peres 11
LODR@onidias-Sepitate er eee ee | SD POTN IS aay eae Ree are eee y
pce onidial>-septate, 17-29 <2 UM kei, eee ok eine ce ele es ete D. septata
CIE C LAG PLCSCll Ltn hee Oe thee No cen ee Os, etwas ae D. novae-guineensis
BUD MS CLACADSENLL, Rie eat ARE. PERNT LS DRC TA TU TERS. WRN Ah) N ke D. eucalypti
ae Ap metac: PLESCH ter ates sterols et tec MRO oy nae eerie on ceeaes eaters D. caatingae
MD et ae ASEM Lee eats Teg hee | ye ean Ue ee T EE D. triseptata
26 ... Cruz &al.
Acknowledgements
Weare deeply indebted to Dr. R.F. Castaneda Ruiz (INIFAT-Cuba) and Dr. Eric McKenzie
(Landcare Research, Auckland-New Zealand) for kindly reviewing the manuscript and
for many suggestions that greatly improved it. Part of the support for this work came
from the Program of Research of Biodiversity in the Brazilian Semi-arid (PPBIO Semi-
arid/Ministry of Technology and Science). The authors L.F.P. Gusmao and A.C.R. Cruz
thank the CNPq and “Programa de Pds-graduacao em Botanica - PPGBot/UEFS”.
A.C.R. Cruz extends thanks to FAPESB (Proc. APR071/2007).
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MYCOTAZXON
Volume 106, pp. 29-40 October-December 2008
Two new species of Polyschema and Vanakripa and other
microfungi recorded from mangrove in Veracruz, Mexico
Rosa MARIA ARIAS MOTA
rosa.arias@inecol.edu.mx
Instituto de Ecologta, A.C., Km. 2.5 antigua carretera a Coatepec 351
Congregacion El Haya, Xalapa, Veracruz, México. C.P. 91070
GABRIELA HEREDIA ABARCA
gabriela.heredia@inecol.edu.mx
Instituto de Ecologia, A.C., Km. 2.5 antigua carretera a Coatepec 351
Congregacion El Haya, Xalapa, Veracruz, México. C.P. 91070
RAFAEL FE. CASTANEDA RUIZ
rfcastaneda@inifat.co.cu
Instituto de Investigaciones Fundamentales en Agricultura Tropical Alejandro de Humboldt
(INIFAT), Calle 1, Esq. 2, Santiago de Las Vegas, C. Habana, Cuba, C.P. 17200
CINTHYA IVONNE BECERRA HERNANDEZ
ivil70@hotmail.com
Instituto de Ecologia, A.C., Km. 2.5 antigua carretera a Coatepec 351
Congregacion El Haya, Xalapa, Veracruz, México. C.P. 91070
Abstract — Two new anamorphic fungi collected on mangrove debris are described
and illustrated. Vanakripa rhizophorae is characterized by variably shaped, globose,
ovoid to ellipsoidal, brown to black, one-celled conidia, and Polyschema nigroseptatum
is distinguished from related species by clavate to ovoid, 2-3(-5)-septate, brown conidia
with black septa. Eleven other species of anamorphic fungi, ten illustrated here, are also
recorded from mangrove.
Key words — systematics, tropical rainforest, conidial fungi
Introduction
Mangrove is an important tropical coast biome; it inhabits a productive
ecosystem able to export energy and materials to adjacent but poor ecosystems.
The mangrove ecosystem that surrounds the La Mancha Lagoon in Veracruz,
Mexico, is characterized by a highly complex landscape containing four
mangrove species: Rhizophora mangle L., Laguncularia racemosa (L.)C.F.
Gaertn., Avicennia germinans (L.)L., and Conocarpus erectus L. (Travieso-Bello
30 ... Arias & al.
& Moreno-Casasola 2006). A distinct group of fungi (manglicolous fungi) occurs
in the mangrove habitat, and mangrove fungi have been reported from a range of
habitats throughout the tropical world (Kohlmeyer 1969, 1980, 1984; Kohlmeyer
& Kohlmeyer 1979, Hyde 1986) and in Malaysia (Jones & Tan 1987). However,
most studies on Mexican mangrove biodiversity have dealt with the flora and
fauna; anamorphic fungi of the region remain poorly explored.
Materials and methods
The mangrove material studied was collected in La Mancha Coastal Research
Center, Veracruz, Mexico (CICOLMA), located 24.5 km north of Cardel
City. Material was air dried for 12 h. Damp chambers were prepared in the
laboratory and incubated during 7-15 days at 28° C. All the species were
studied directly from naturally occurring plant debris. Slides were mounted
in polyvinyl alcohol. Specimens cited have been deposited at the herbarium in
Xalapa (XAL), Veracruz, Mexico. }
Taxonomy
Polyschema nigroseptatum R.M. Arias, R.F. Castafieda & Heredia, anam. sp. nov.
MycoBank, MB 511869 Figs. 3-7
COLONIE in substrato naturali effusae, nigrae. MyCcELIuM partim superficiale et partim
in substrato immersum. Hyphae septatae, ramosae, laeves, brunneae. CONIDIOPHORA
conspicua, erecta, moniliformia, laevia, 20-38 x 5-7 um, brunnea. CELLULAE
CONIDIOGENAE monotreticae, globosae vel clavatae, determinatae, 8-10 x 5-7 um, pallide
brunneae, laeves. CONIDIA Solitaria, acrogena, clavata usque ad ovoidea, 2-3(-5)-septata,
41-45 x 19-22 um, brunnea, atrobrunnea vel nigra ad septa, laevia, sicca.
Hototyree: MEXICO. VERACRUZ: La Mancha Coasta Research Center, on bark of
Rhizophora mangle, 2.VIII.1995, G. Heredia (HOLOTYPE: XAL CB896).
ETtyMoLoGcy: Latin, nigroseptatum, referring to the dark brown to black septa.
COLONIES on the natural substrate effuse, black. MyceLium superficial and
immersed. Hyphae septate, branched, smooth-walled, brown. CONIDIOPHORES
conspicuous, erect, moniliform, smooth-walled, 20-38 x 5-7 um, brown.
CONIDIOGENOUS CELLS monotretic, globose or clavate, determinate, 8-10 x
7-10 um, pale brown, smooth-walled. Conip1a solitary, acrogenous, clavate to
ovoid, 2—3(-5)-septate, 41-45 x 19-22 um, brown, but dark brown to black at
the septa, smooth-walled, dry.
Notes: Sixteen species are recognized in the genus Polyschema H.P. Upadhyay
(Castaneda et al 2000): P. bicellulare Shearer, P chambalense I.J. Joshi et al.,
Figs 1-6. 1-2. Vanakripa rhizophorae (from XAL CB895). 1. Conidia, separating cells and
conidiogenous cells. 2. Sporodochium. 3-6. Polyschema nigroseptatum (from XAL CB896).
3-4. Conidiogenous cells and conidia 5-6. Conidiophores, conidiogenous cells and conidia. Scales
are indicated by bars.
Polyschema & Vanakripa spp. nov. (Mexico) ... al
oe
a
oot
seve,
Sou
pod
32 ... Arias & al.
Fic. 7. Polyschema nigroseptatum (from XAL CB896).
Conidiophore, conidiogenous cells and conidia. Scales are indicated by bars.
P. clavulatum (Cooke & Harkn.) M.B. Ellis, P congolense Reisinger & Kiffer,
P. cubense Mercado & J. Mena, P. indicum Behera et al., P. larviforme (Fairm.)
M.B. Ellis, P lignicola V.G. Rao & K.A. Reddy, P. obclaviforme R.F. Castafieda &
Decock, P. olivaceum (Ellis & Everh.) M.B. Ellis, P queenslandicum Matsush., .
P. sagari J.P.N. Pandeya & S.B. Saksena, P. terricola H.P. Upadhyay, P. variabile
D.P. Tiwari et al., P. venustum R.F. Castafieda et al., and P yakuense Matsush.
(Most authors have treated the genus Polyschema — based on the Greek neuter
noun, oyna — as feminine instead of neuter; all the appropriate epithet gender
corrections have been incorporated here.) Polyschema toruloides Matsush. was
not included in Castafieda et al. (2000), since this species was described from
Polyschema & Vanakripa spp. nov. (Mexico) ... 33
V8jA cultures, and the conidiogenous cells are multilocal, globose to broadly
obovate, black or dark brown, verrucose. These characters relate P. toruloides
more closely to the genus Dwayabeeja Subram., but we do not propose a new
combination here. Polyschema indicum D.P. Tiwari & P.D. Agrawal (published
June 1973), which is an illegitimate later homonym of P. indicum Behera et al.
(published March 1973; see Commonwealth Mycological Institute 1975, Index
Fungorum 2008), was also not included in Castaneda et al. (2000). Tiwari &
Agrawal (1973) described their species from on potato dextrose agar cultures
and the conidia are variably shaped: subglobose, globose, obovate, oval, clavate,
obpyriform to pyriform, (1-)2-4-septate (mostly 2-septate), 12.7-29.4 x
8.3-13.3 um. In fact, the conidia of P indicum D.P. Tiwari & P.D. Agrawal
strongly resemble those of P indicum Behera et al. Polyschema nigroseptatum
is superficially close to P. variabile in the shape of the conidia, but the latter has
smaller conidia (9.0-28.5 x 6.0-11.5 um).
Key to Polyschema species
ram@onidiasimooth-walled, (> onesie: ome eee meer eee, RIES 2. 2
Epa onidiacwithfornariented, Wall Sumer ees tae earn ert ee cesta Siem, SER i
pameonidia with one septiin = ane Merwe. | ewe Me TT) eee eer ae 3
Do Gonidia with more: than Onesepiliniees aes eat, meme tens cress setts e), Sent 28, 4
3a. Conidia 14-21 x 7-12 um, obovoid or pyriform, pale to mid brown, with a dark
brown band at the septum; apical cell longer than the basal cell ..... P. lignicola
3b. Conidia 10-12 x 5-6 um, subclavate to obovoid, reddish brown, without a dark
brown band at the septum and with the apical cell three times longer than the
basal cellapeere Se raster Sent ree ee ee a PC ST ES P. bicellulare
4a. Conidia 2-3-septate, oblong, clavate or ellipsoidal, reddish brown, darker at the
SEDiaw lo 20 2g Oy [Tear re age eens eter a gs tate es eee Sc P. cubense
EDR OLIGTA Willies OliIOLe GCP tat saan ee ma ern eect Ae amt ee ee ee ee eee 5
5a. Conidia 30-48 x 6-7 um, obclavate, (4—)5-septate, brown at the middle and pale at
ACS AYO RP a ee Lie eve yeni cee Wie ge Rt ie Sede daira ie A ie ae P. obclaviforme
Pomc onidia wider tual LOMUT eS —S-SCDlAlC mg. Metre ee re etn te, 6
6a. Conidia 30-46 x 12-16 um, clavate, 3-5-septate, pale to dark reddish brown, with
basabeell(palerthan the othergeraees.. Meee pee en P. clavulatum
6b. Conidia 41-45 x 19-22 um, clavate, ovoid, 2-3(-5)-septate, dark brown to black at
dT aOLC TY Cee BEEN WA re h VES. tt ey em gee Sees P. nigroseptatum
Pam Olli cliatverTucOScmtisse ts een Ria) Angee 4 00 yas ce hres ees We RP eae ea ee NE 8
Bec oniciastuberculate Om pap atc eet. eos Ae AP cere ern ete 7 gee ir
8a. Conidia with oblique septa, mostly subglobose, pyriform or clavate, mid to dark
DIOW al 3 eS 1 8 LT ret cr ae ek Sie oe P. terricola
Bee OniGia Ww TLDOULODIGUEG:SEDtdg aitah 6 che ete ae a eek et heen Pe 9
34 ... Arias & al.
9a: Conidia usually withasmaximum O87 septa sri eer ye eh ee ae ae 10
9b: Conidia withwmorethansasep tags.) nes See era eee ee eee 14
10a? Conidia ellipsoidalt0clavate.or pyritora m1. eee ees ee one Oar etre ik 11
10b}Conidia cylindrical iconstricted at the septinp a ee ereeee Sarton tie eae 13
11a. Conidia 0-5-septate, clavate, gray-brown or dark brown at the upper two cells,
pale brown towards the base, 9.0-28.5 x 6.0-11.5um .............. P. variabile
lib Conidia‘usually, llipsoidtompyriforings ses se eee hee tana ee 12
12a. Conidia 1-3(mostly 2)-septate, ellipsoidal, sometimes clavate, pale golden brown,
L6=26°58 8-0 ee eee oe rn. eee eee ere ee ane P. congolense
12b. Conidia 2—4-septate, ellipsoidal or pyriform, mid dark golden brown, paler
atithe.ends>15230'< LOSE fing eee ere. OAM ee mere ed, en P. indicum
13a. Conidia 2-7-septate, constricted at the septa, mid golden brown, 17-37 x
GS inne! Alan hie nemels 09, SES Ry Ne Aes ALON ee ee Aree A: P. olivaceum
13b. Conidia 1-5-septate, constricted at the septa, pale brown, 5-20 x 3-8 um
RE Pe ae I oo, PPO ie Oe Oe ee Gee Sens ek SACaIt
14a. Conidia sometimes catenate, clavate, obclavate, cylindrical to sigmoid, pale
brown,.3=9-septate,.13=19"46- lili, -atee. re ent P. chambalense
14b. Conidia not catenate, clavate, obclavate or ellipsoidal, reddish brown to dark
reddish brown, 4-12-septate, 30-80 x12 -20 um ................ P. larviforme
15a. Conidia tuberculate, navicular or broadly fusiform, dark brown or black,
brown at the ends, 5-10-septate, 30-60 x 13-20 um .............. P. venustum
15b-Conidia:papllatem 2. ee ta). LSA Pep 2 a ede he ie 16
16a. Conidia ellipsoidal or oblong-ellipsoidal, 2—3-septate, dark brown, paler
towards:the endsyl3=22)< 6 Simi ete pee pepe See ot, ae ere P. yakuense
16b. Conidia clavate to ellipsoid, central cells brown, pale brown at the ends,
2-4(mostly 3)-septate, 20-36 x 9-13 um ................-- P. queenslandicum
Vanakripa rhizophorae R.M. Arias, Heredia & R.E. Castafieda anam. sp. nov.
MycoBank, MB 511867 Figsalo2.6
CoNIDIOMATA sporodochialia, superficialia, dissita, pulvinata, nigra. MYCELIUM partim
superficiale et partim in substrato immersum, ex hyphis septatis, ramosis, hyalinis, laevibus
compositum. CONIDIOPHORA inconspicua, mononematosa, hyalina, interdum in cellulis
conidiogenis reducta. CELLULAE CONIDIOGENAE hologenosae, determinatae, cylindricae,
hyalinae, in conidiophoris incorporatae. CELLULAE SEPARANTES Clavatae, usque ad
vermiformes, 29.4-53.9 x 3.9-9.3, multiguttulatae, laeves, hyalinae. ConrpIA solitaria,
acrogena, variabilia, globosa, ovoidea usque ad ellipsoideae, unicellularia, 16.6-22.3 x
6.3-14.7 um, atrobrunnea usque ad nigra.
HoLotyPe: MEXICO. VERACRUZ, La Mancha Coasta Research Center, on bark of
Rhizophora mangle, 2.V11I.1995, G. Heredia (HOLOTYPE: XAL CB895).
EryMo_oGcy: Latin, rhizophorae, refers to its growing on Rhizophora mangle host.
COLONIES on the natural substrate effuse, black. Conipiomata sporodochial,
scattered, pulvinate. MYCELIUM superficial and immersed composed of septate,
Polyschema & Vanakripa spp. noy. (Mexico) ... 35
Fig. 8. Vanakripa rhizophorae (from XAL CB895).
Conidia and separating cells. Scales are indicated by bars.
branched, hyaline, smooth-walled hyphae. CoNrDIOPHORES undifferentiated,
mononematous, hyaline, sometimes reduced to conidiogenous cells.
CONIDIOGENOUS. CELLS hologenous, determinate, cylindrical, hyaline,
integrated. SEPARATING CELLS clavate to vermiform, 29.4-53.9 x 3.9-9.3 um,
multi-guttulate, smooth-walled, hyaline. Conip1a solitary, acrogenous, variable
in shape, globose, ovoid to ellipsoidal, 16.6-22.3 x 6.3-14.7 um, aseptate,
smooth, brown to black.
Notes: The genus Vanakripa Bhat et al. currently comprises five species:
V. ellipsoidea K.M. Tsui et al., V. fasciata R.F. Castafieda et al., V. gigaspora Bhat
36 ... Arias & al.
et al., V. minutiellipsoidea Pinnoi and V. parva Bhat et al. Vanakripa rhizophorae
produces black conidia with hyaline, clavate separating cells, but the conidia
are variable in shape, ranging from globose, ovoid to ellipsoidal; it is close to V.
parva but differs in having larger conidia.
Key to Vanakripa species
la: Conidia one=celledigrs ys re eerie lls hls cs ey, 2
1b. Conidia with a submedian septum, clavate to obpyriform, smooth-walled, dark
brown to black, basal cell pale brown, 33-40 x 19-23 um.......... V. gigaspora
2a. Conidia small, 7-10 x 4-6 um, ovoid, with rounded apex, brown to dark brown,
STHOOCHS WalleCl Rt Mes se Rte etree actennes other hPa ey tea oN eh ae en en ee V. parva
2b, Conidia muchJarcersmoreithan) 20941 4 ini ee oe eats Cee ee ae 33
3a. Conidial shape variable, globose, ovoid to ellipsoidal, 16.6-22.3 x 6.3-14.7 um,
smooth=walledidarkibrownato Dlac ka eee re ee V. rhizophorae
3b. Conidial' shape uniform | aegis ee RS Ee HE ea iF 4
4a. Separating cells relatively narrow, 17-30 x 6-9 um, clavate, hyaline; conidia
ellipsoidal, with rounded apex, brown to dark brown with a central zone or a
black transverse band, smooth-walled, 24-34 x 16-23 um........... V. fasciata
Ab. Separating cellsimuch wider more thanel0 (impees aa ee cae sn ae ee ee 5
5a. Conidia 25-33 x 16-23 um, ellipsoidal, with rounded apex and base, dark
brown to black, smooth-walled; separating cells clavate to vermiform,
30270610222! aim, hyaline ea ase oo ene ae V. minutiellipsoidea
5b. Conidia longer; 33-42 x 20-25 um, ellipsoid, with rounded apex, dark brown
to black, smooth-walled, separating cell clavate to vermiform, 30-72 x
Li1-4] 6 Lims.smoothawalled) hyalincee stent cern eee V. ellipsoidea
Other anamorphic fungi recorded from mangrove
Coniosporium memorandum (Penz. & Sacc.) M.B. Ellis, Mycol. Pap. 125: 2 (1971).
(Fic. 16)
SPECIMEN EXAMINED: MEXICO.. VERACRUZ, La Mancha Coasta Research Center, on
dead leaves of Avicennia germinans, 2.VII.1995, G. Heredia (CB 884, CB884-1).
Corynespora kamatii (V.G. Rao) M.B. Ellis, More Dematiaceous Hyphomycetes
(Kew): 376 (1976). (Fic. 11)
SPECIMEN EXAMINED: MEXICO. VERACRUZ, La Mancha Coasta Research Center, on
dead stems of Rhizophora mangle, 11.VIII.1995, G. Heredia (CB885, CB885-1).
Fics 9-18. 9 a-b. Virgaria nigra, a - Conidiophores b - Conidia. 10. Periconia minutissima,
conidiophores, condiogenous cells and conidia. 11. Corynespora kamatii, conidium. 12.
Ellisembia adscendens, conidium. 13. Helicoma proliferens, conidiogenous cell and conidium. 14.
Intercalarispora nigra, conidium. 15. Mycoenterolobium platysporum, conidium. 16, Coniosporium
memorandum, conidia. 17. Pseudopetrakia kambakkamensis, conidium. 18. Tetracoccosporium
aerium, conidia. Scales are indicated by bars.
Polyschema & Vanakripa spp. nov. (Mexico) ... 37
wd
n OZ
"SUITE RC EET EPEC PLOREMDCTONECRE OPEL
nin Sz
38 ... Arias & al.
Ellisembia adscendens (Berk.) Subram., Proc. Indian Natl. Sci. Acad., B, Biol. Sci.
58(4): 183 (1992). (Figs 12)
SPECIMEN EXAMINED: MEXICO. VERACRUZ, La Mancha Coasta Research Center,
on dead stems of Avicennia germinans, 1.VUI.1995, G. Heredia (CB886, CB886-1,
CB886-2).
Helicoma proliferens Linder, Ann. Missouri Bot. Gard. 16: 309 (1929). (Fic. 13)
SPECIMEN EXAMINED: MEXICO. VERACRUZ, La Mancha Coasta Research Center, on
bark of Rhizophora AGES 1.VIII.1995, G. Heredia (CB887).
Intercalarispora nigra J.L. Crane & Schokn., in Schoknecht & Crane, Canad. J. Bot.
61(8): 2243 (1983). (Fic. 14)
SPECIMEN EXAMINED: MEXICO. VERACRUZ, La Mancha Coasta Research Center, on
bark of Rhizophora mangle, 2.V1I.1995, G. Heredia (CB894).
Mycoenterolobium platysporum Goos, Mycologia 62(1): 172 (1970). (Fic. 15)
SPECIMEN EXAMINED: MEXICO. VERACRUZ, La Mancha Coasta Research Center, on
stalk of Rhizophora mangle, 17.VIII.1995, G. Heredia (CB888, CB888-1).
Periconia minutissima Corda, Icon. fung. (Prague) 1: 19 (1837). (Fic. 10)
SPECIMEN EXAMINED: MEXICO. VERACRUZ, La Mancha Coasta Research Center, on
stalk of Rhizophora mangle, 2.VUI.1995, G. Heredia (C 889).
Pseudopetrakia kambakkamensis (Subram.) M.B. Ellis, Mycol. Pap. 125: 4 (1971).
(Fic. 17)
SPECIMEN EXAMINED: MEXICO. VERACRUZ, La Mancha Coasta Research Center, on
bark of Rhizophora mangle, 1.V1I.1995, G. Heredia (CB890).
Tetracoccosporium aerium P.C. Misra & P. Srivast., Mycotaxon 4(1): 276 (1976).
(Fic.18)
SPECIMEN EXAMINED: MEXICO. VERACRUZ, La Mancha Coasta Research Center, on
bark of Avicennia germinans, 23.VIII.1995, G. Heredia (CB891, CB891-1).
Verticillium chlamydosporium var. catenulatum (Kamyschko ex Onions & G.L.
Barron) W. Gams, Netherlands. J. Pl. Pathol. 94(3): 134 (1988).
SPECIMEN EXAMINED: MEXICO. VERACRUZ, La Mancha Coasta Research Center, on
bark of Rhizophora mangle, 2.V1II.1995, G. Heredia (CB892, CB892-1).
Virgaria nigra (Link) Nees, Syst. Pilze: 54 (1817). (FiG.9)
SPECIMEN EXAMINED: MEXICO. VERACRUZ, La Mancha Coasta Research Center, on
bark of Avicennia germinans, 1.VII.1995, G. Heredia (CB893).
Polyschema & Vanakripa spp. nov. (Mexico) ... 39
Acknowledgements
We are deeply indebted to Lori M. Carris (Washington State University) and U. Braun
(Martin-Luther- Universitat) for kindly reviewing the manuscript. We are grateful to the
CYTED Program (REDEMIC JXII) for funding the scientific collaboration among the
authors and the Cuban Ministry of Agriculture for facilities. The author RFCR thanks
Drs. David W. Minter, Pedro Crous, Uwe Braun, Ludmila Marvanova, Cony Decock,
Jerry A. Cooper for their generous and valuable assistance with literature not available
in our library.
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MYCOTAXON
Volume 106, pp. 41-63 October-December 2008
New cercosporoid fungi from the Brazilian Cerrado
1. Species on hosts of the families Anacardiaceae,
Araliaceae, Bombacaceae, Burseraceae and Celastraceae
ANTONIO HERNANDEZ-GUTIERREZ”” & JOSE C. DIANESE”
*icarmine@unb. br
‘Instituto de Ciéncias Biologicas. Campus Universitario do Guamd
Universidade Federal do Para, Rua Augusto Corréa. 66.075-110 Belém, PA, Brazil
?Departamento de Fitopatologia, Universidade de Brasilia
70.910-900 Brasilia, DE, Brazil
Abstract — New cercosporoid hyphomycetes are described from the Brazilian
Cerrado on native plants of the families Anacardiaceae (Pseudocercospora astronii,
P. astroniiphila), Araliaceae (Passalora schefflerae), Bombacaceae (Cercospora ochromae,
Pseudocercospora eriothecae, P. pseudobombacis), Burseraceae (Pseudocercospora protii),
and Celastraceae (Prathigada austroplenckiae, Pseudocercospora austroplenckiae).
Key words — fungal taxonomy, mycodiversity, Mycosphaerellaceae, tropical
hyphomycetes
Introduction
The Cerrado is the second largest Brazilian biome, surpassed only by the
Amazonian Forest. ‘The floristic richness of the Cerrado includes a total of 6671
native taxa, distributed in 170 families and 1144 genera. There are 6429 species
and 451 varieties and subspecies, respectively, including 264 species belonging
to the pteridophytes (in 19 families and 51 genera), two to the gymnosperms
(Podocarpus, Podocarpaceae) and 6060 to the angiosperms (in 150 families and
1092 genera) [according to Mendonga et al. (1998)].
Although the majority of Cerrado plants are well documented, little is known
about their associated microfungi. The exploration of Cerrado microfungi dates
back to the end of the 19th century with first contributions by some American
and European mycologists, including P. Hennings, E.W.D. Holway and HLS.
Jackson. However, the most important contributions were made by the Brazilian
mycologists A.P. Viégas and A.C. Batista, who described more than 240 species
‘Portion of the Doctor’s Thesis of the senior author
42 ... Hernandez-Gutiérrez & Dianese
from the Cerrado (Dianese et al. 1997). The most comprehensive contribution
to the knowledge of the cercosporoid fungi of Brazil was undoubtedly published
by Viégas (1945), which was later used by Chupp (1954) for the preparation of
his monograph of Cercospora Fresen. Since then, only fragmentary treatments of
Brazilian cercosporoid fungi were published (Dianese & Camara 1994, Medeiros
1994, Barreto & Evans 1995, Crous & Braun 1996, Inacio et al. 1996, Crous et
al. 1997, Inacio & Dianese 1999, Furlanetto & Dianese 1999, Braun et al. 1999,
Braun & Freire 2002, 2004, 2006). In this paper cercosporoid hyphomycetes
found on native Cerrado plants belonging to five host families are described
following the generic concepts introduced by Deighton (1967, 1974, 1976)
and updated by Braun (1999) as well as Crous & Braun (2003). The collections
studied are deposited in the Mycological Collection of the Herbarium of the
University of Brasilia (Herbarium UB - mycological collection).
Materials and methods
Starting in 1992, Cerrado fungi were extensively collected from different regions
representative for this biome (Dianese et al. 1997). The plant samples containing the
fungi were pressed and dried in a drying-chamber during 3-4 days at ca. 50°C, then they
were exsiccated, labeled and frozen for 7-8 days at temperatures between -—30°C and
—50°C, and finally they were re-dried in an oven during 24 h at temperatures between
50-70°C before being deposited in the Mycological Collection of the Herbarium
UB. For the study of cercosporoid fungi associated with different hosts, observations
under a stereomicroscope were made to describe the lesions as well as the location and
characteristics of the fungal colonies. For the morphological studies, each specimen
was mounted on slides using the following media: lacto-glycerol/cotton blue, glycerol-
KOH/basic floxine and polyvinyl alcohol (PVA 4.165 g, acetic acid 25 ml, glycerol 2.5
ml, distilled water 25 ml; this preparation was autoclaved for 15 min. and left to rest
for 24 h before use) (Omar et al. 1979). Mounting media without stains were also used
to determine the exact color of different structures, and cotton blue and basic floxine
solutions were used to observe the conidial septation and conidiogenous cells. To verify
the relationship between the fungus and the host, pieces of the dried vegetal material
containing the lesions were re-hydrated for 4-12 h using an aqueous solution containing
10% ethanol and 0,1% Tween-20. Then 7-10 um thick sections were produced using a
freezing microtome (MICRON, HM505E, Micron Laborgerate, Walldorf, Germany).
Fungal material was studied using a compound light microscope (Zeiss Ultraphot
III, Carl Zeiss, Oberkochen, Germany) provided with a camera lucida for drawings.
A calibrated ocular micrometer was used to measure (length and width, or diameter) _
fungal structures (stromata, conidiophores, conidiogenous cells and conidia). Whenever
possible between 40 and 50 measurements were done. Numerical data such as number
of conidiophores per fascicles, septa and conidiogenous loci, were observed.
A detailed drawing of each specimen was produced showing the location of lesions,
extension and shape of the symptoms on the host, and colony features, together with
a camera lucida scaled drawing of a micrometric section of the infected plant tissue
Cercosporoids from the Brazilian Cerrado ... 43
containing details of stromata, conidiogenous cells and conidia. The drawings were
prepared using A4 white layout paper using Pilot (Japan) pens, with points of 0.1, 0.2,
0.3, and 0.5 mm diam.
Taxonomy
Cercosporoid fungi on Anacardiaceae
The family Anacardiaceae comprises 76 genera and 600 species, tropically
and sub-tropically distributed, except for the genus Rhus occurring in
temperate regions. This family is represented in the Cerrado with eight
genera (Anacardium, Astronium, Lithrea, Myracrodruon, Schinopsis, Schinus,
Spondias and Tapirira) and 20 species, among them five species belonging in
Astronium (Mendonga et al. 1998). Numerous cercosporoid species have been
described from hosts of the Anacardiaceae (Viegas 1945, 1961; Chupp 1954,
Braun 1995, 1998; Hsieh & Goh 1995). On Astronium spp. two cercosporoids
are known: Passalora myracrodruonis (Inacio & Dianese) U. Braun & Crous
(=Mycovellosiella myracrodruonis Inacio & Dianese), describedon Myracrodruon
urundeuva M. Allemao (= Astronium urundeuva (M. Allemao) Engl.) from the
Brazilian Cerrado (Inacio & Dianese 1999), and Pseudocercospora astroniicola
U. Braun & E O. Freire (Braun & Freire 2002), introduduced for a cercosporoid
fungus on Astronium fraxinifolium Schott in the Brazilian State of Ceara. Two
species belonging in other fungal groups are also known on A. fraxinifolium,
an important Anacardiaceae from the Cerrado, viz. Phyllachora astronii Speg.
(Medeiros 1994) and the rust fungus Kimuromyces cerradensis Dianese et al.
(Dianese et al. 1995).
Two new Pseudocercospora Speg. species on A. fraxinifolium and
A. urundeuva have recently been found in the Cerrado.
Pseudocercospora astronii A. Hern.-Gut. & Dianese sp. nov. FIG. 1
MycoBank, MB 512026
Pseudocercosporae astroniicolae similis, sed hyphis superficialibus et conidiophoris
solitariis nullis, cellulis conidiogenis geniculatis, locis conidiogenis subconspicuis, leniter
prominentibus, planatis, leviter incrassatis, conidiis longioribus, 20-76 um, deinde 4-7-
septatis, septo basali constricto et incrassato.
SPECIMENS EXAMINED: BRAZIL. MINAs GERAIS: DIVINOPOLIS, Feira Agropecuaria, on
living leaves of Astronium fraxinifolium (Anacardiaceae), 12 Jun 1995, leg. C. A. Indcio
132, holotype (UB Col. Micol. 8992). Minas GERAIS: DIVINOPOLIS, Feira Agropecuaria,
on living leaves of Astronium urundeuva (Anacardiaceae), 26 Jun 1995, leg. C. A. Indcio
318, paratype (UB Col. Micol. 17811).
Lesions 1-14 mm diam., irregular, amphigenous, dark brown, coalescent,
without a definite margin. COLONIES mainly hypophyllous. Srromata brown to
dark brown, subepidermal, textura globosa, 28-65 um diam. CONIDIOPHORES
fasciculate, straight to slightly curved, lageniform, short, 17-24 um long, 3-4 um
44 ... Hernandez-Gutiérrez & Dianese
Fig. 1. Pseudocercospora astronii on Astronium fraxinifolium. Distribution of lesions
on a leaf (top left); stroma and conidiophores with conidiogenous cells bearing
inconspicuous conidiogenous loci sided by obclavate conidia; a detailed view of
the conidiogenous cell (right). Based on the holotype (UB Mycol. Col. 8992).
diam. in the widest part, light brown to olivaceous, paler toward the apex, 0-1-.
septate, smooth, apex truncate, 1-3 geniculate, thin-walled. CONIDIOGENOUS
CELLS integrated, terminal, polyblastic, sympodial, geniculate; conidiogenous
loci slightly prominent, smooth, barely thickened, but not pigmented. CoNIDIA
solitary, straight, curved or flexuous, subcylindrical to narrowly obclavate,
aseptate when young, 4—7-septate when mature, basal septum constricted and
Cercosporoids from the Brazilian Cerrado ... 45
thicker than the others, 20-76 um long, 2—4 um diam. at the widest part, 1-2
um diam. at the base and 1-3 um diam. close to the apex, mostly smooth,
sometimes finely verruculose.
Pseudocercospora astroniiphila A. Hern.-Gut. & Dianese sp. nov. Fic. 2
MycoBank, MB 512035
Pseudocercosporae astronii similis, sed conidiophoris saepe cylindraceis, 1-3-septatis et non
geniculatis, conidiis saepe rectis, levibus et uniformiter distincte septatis, sine constrictione.
Differt a P. astroniicola hyphis superficialibus et conidiophoris solitariis nullis.
SPECIMENS EXAMINED: BRAZIL. MINAS GERAIS: DIVINOPOLIS, Feira Agropecuaria, on
living leaves of Astronium urundeuva (Anacardiaceae), 26 Jun 1995, leg. C. A. Indcio
320, holotype (UB Col. Micol. 8999). Minas GERAIS: DIVINOPOLIS, Feira Agropecuaria,
on living leaves of Astronium urundeuva (Anacardiaceae), 1 Jul 1995, leg. C. A. Indcio
314, paratype (UB Col. Micol. 8992).
Lesions 2-5 mm diam., irregular, brown. Mycelium immersed in the
substratum. CoLontes hypophyllous. Stromata 18-60 um diam., well-
developed, textura globosa, subcuticular or subepidermal, erumpent, olivaceous.
CONIDIOPHORES compactly fasciculate, straight, flexuous or curved, cylindrical
to slightly lageniform, 2—3-septate, 15-21 um long, 3-4 um diam. at the widest
part, olivaceous, thin-walled, smooth. CONIDIOGENOUS CELLS integrate,
terminal, mostly monoblastic, occasionally sympodial; conidiogenous loci
inconspicuous, flattened, non-pigmented. Conip14 solitary, straight, curved or
flexuous, cylindrical to slightly obclavate, 0-6-septate, 28-72 um long, 2-4 um
diam. at the widest part, 1-2 um diam. at the base and at the apex, rounded
at the apex; conidial base truncate to obconical truncate, light olivaceous to
subhyaline, smooth, unthickened.
COMMENTS: Pseudocercospora astroniicola (Braun & Freire 2002), described
from the Brazilian State of Ceara on A. fraxinifolium, is the only comparable
species on a host belonging to the genus Astronium including the closely allied
genus Myracrodruon. The two new species (P. astronii and P. astroniiphila)
are distinguished from P astroniicola by having uniformly fasciculate
conidiophores, i.e. superficial hyphae with solitary conidiophores are lacking.
In addition, P astronii differs from P astroniicola in having conidiogenous
cells with subconspicuous, slightly prominent, truncate conidiogenous loci
and longer conidia, 4—7-septate when mature, constricted at the basal septum,
which is thicker than the other septa. Pseudocercospora astronii was found on
A. fraxinifolium as well as M. urundeuva, on the latter host sometimes together
with P. astroniiphila on the same lesion, but P. astroniiphila was confined to
leaves of A. urundeuva. Pseudocercospora astronii differs from P. astroniiphila
in having strictly lageniform, geniculate, l-septate conidiophores and
straight or sometimes slightly curved and slightly verruculose conidia, with
46 ... Hernandez-Gutiérrez & Dianese
ronson
“et
sernscagcnneenponnenngnenees : * CS ae
> i setae MN ATION
* 5 eee
lee as Soanadnnisicnrtion eo
Se eaten ee Man roa a
f = 4
; e £ §
;
et
Herre ~
Fig. 2. Pseudocercospora astroniphila on Astronium urundeuva. Lesions on a leaf (top
right); erumpent-subcuticular stroma with a fascicle of conidiophores bearing
conidiogenous cells; a representative sample of conidia; a detailed view of a
conidiophore (bottom right). Based on the holotype (UB Mycol. Col. 8999).
a constriction at the thicker basal septum (versus conidiophores cylindrical,
2-3-septate, seldom geniculate and conidia generally straight, smooth-walled,
with conidial septa conspicuous and without constrictions in P. astroniiphila).
Pseudocercospora comocladiae (Petr. & Cif.) Deighton and P mombin (Petr.
& Cif.) Deighton are two morphologically similar species described on hosts
belonging to other genera of the Anacardiaceae (Chupp 1954, Deighton 1976).
P. comocladiae is distinct from P. astronii and P. astroniiphila by its very short,
aseptate conidiophores, 5-15 um long. The conidia are uniformly septate,
without any constrictions. P mombin differs from P. astronii in having aseptate
conidiophores and uniformly septate conidia without any constrictions, and
from P. astroniphiila in shorter, aseptate conidiophores. P. rhinocarpi U. Braun
Cercosporoids from the Brazilian Cerrado ... 47
& Crous (Crous & Braun 2003) on Anacardium in Venezuela has much smaller
stromata, only 10-30 um diam., and shorter, 1—-4-septate conidia, 10-50 x 1.5-3.5
um in comparison to both new species.
Cercosporoid fungi on Araliaceae
Seventy genera belong to the family Araliaceae, distributed mainly in
three tropical zones: the Indo-Malayan region, Australia and Neotropica. Six
genera are known from the Cerrado: Dendropanax, Didymopanax, Oreopanax,
Pentapanax, Sciadodendron and _ Schefflera, comprising sixteen species
(Mendonga et al. 1998). Cercosporoid hyphomycetes in the genera Cercospora,
Cercosporella Sacc., Pseudocercospora, and Pseudocercosporella Deighton have
been described from several hosts of the Araliaceae (Chupp 1954, Viégas 1961,
Farr et al. 1989, Hsieh & Goh 1990, Guo & Hsieh 1995, Braun 1995, 1998, Crous
& Braun 2003). However, the new species found in the Cerrado on Schefflera
macrocarpa (Cham. & Schltdl.) Frodin represents the first Passalora Fr. species
on an Araliaceae. There is no comparable species.
Passalora schefflerae A. Hern.-Gut. & Dianese, sp. nov. FIG. 3
MycoBank, MB 512037
Laesiones minutae, punctiformes vel 7-8 mm diam., amphigenae, angulares vel
irregulares, pallido- vel atro-brunneae. Coloniae abaxiales, punctiformes, 7-8 um diam.,
velutinae, inter trichomata. Mycelium immersum. Stromata 25-96 um diam., primo
substomatalia, deinde suprastomatalia, e textura globosa composita, bene evoluta, atro-
brunnea. Conidiophora in fasciculis laxe aggregatis, ramosa, inter trichomata. Cellulae
conidiogenae in conidiophoris incorporatae, terminales vel intercalares, polyblasticae,
cicatricibus conspicuis, incrassatis et fuscatis, 2-4 um latis. Conidia solitaria, recta,
curvata vel flexuosa, cylindrica, leviter clavata vel obclavata, ad basem obconica vel
obconico-truncata, pallido-brunnea vel olivacea, laevia, 21-96 wm longa, 5-10 wm lata in
parte latissima, 2-6 um lata ad basem, 3-10 um lata prope apicem, 0—7-septata, septo in
medio crassiori et atriori; hilum incrassatum et fuscatum.
SPECIMENS EXAMINED: BRAZIL. Gots: CRISTALINA, FAZENDA Nova INpIA, on living
leaves of Schefflera macrocarpa (Araliaceae), 19 Apr 1993, leg. J.C. Dianese 1058, holotype
(UB mycol. col. 4464). Disrriro FEDERAL: BRASILIA, Parque Nacional de Brasilia, on
living leaves of S. macrocarpa (Araliaceae), 22 Jan 1998, leg. E.A. Teixeir 6, paratype
(UB mycol. col. 15831). GorAs: CRISTALINA, Fazenda Nova India, on living leaves of
Schefflera macrocarpa (Araliaceae), 13 Oct 1993, leg. J.C. Dianese 1316, paratype (UB
mycol. col. 5155).
LEsIONS small, punctiform or up to 7-8 mm diam., amphigenous, angular or
irregular, light to dark brown. CoLontgs abaxial, velvety. MYCELIUM immersed.
STROMATA at first substomatal, later suprastomatal-erumpent, dark brown, well-
developed, textura globosa, 25-96 um diam. CONIDIOPHORES in loose fascicles,
branched, intermixed with the trichomes. CONIDIOGENOUS CELLS integrated,
terminal or intercalary, polyblastic; conidiogenous loci protuberant, thickened,
darkened, 2-4 um diam. Conip1A solitary, 21-96 um long, 5-10 um diam.
48 ... Hernandez-Gutiérrez & Dianese
Fig. 3. Passalora schefflerae on Schefflera macrocarpa. Lesions on a leaf (top right);
a representative sample of conidia; branched superficial secondary mycelium;
substomatal erumpent stroma bearing a fascicle of conidiophores. Based on the
holotype (UB Mycol. Col. 4464).
at the widest part, 2-6 um at the base and 3-10 um near the apex, simple,
straight, curved or flexuous, light-olivaceous to olivaceous, smooth, cylindrical
to slightly clavate or obclavate, obconically truncate at the base, 0-7-septate,
with a thicker, darker septum in the middle; hilum thickened and darkened.
COMMENTS: Due to the basic features of this cercosporoid fungus on Schefflera,
i.e. conspicuously thickened and darkened conidiogenous loci and conidial hila
Cercosporoids from the Brazilian Cerrado ... 49
as well as pigmented conidia, it can be assigned to Passalora as emended by
Crous & Braun (2003). Nevertheless, this species is very unusual by having very
large stromata with densely fasciculate, frequently branched conidiophores
which are intermingled among trichomes. However, characteristics of the
stromata within the Cercospora complex are considered “...useful for species
delimitation, but inappropriate for generic discrimination...” (Braun 1995).
Branched appendages are unusual in Passalora, but they may occur in a few
species, e.g. P. bacilligera (Mont. & Fr.) Mont. & Fr. (Ellis 1971). However, the
intensity and frequency of branching in P. schefflerae is stronger than in all
comparable species.
Cercosporoid fungi on Bombacaceae
Bombacaceae is a pantropical family with 28 genera, distributed mainly in
Neotropica. In the Cerrado, this biome encompasses seven genera (Bombax,
Cavanillesia, Ceiba, Chorisia, Eriotheca, Pseudobombax and Quararibea) and
17 species, among them five belonging in Eriotheca (Mendonga et al. 1998).
Ochroma comprises bombacaceous species typical of the Amazonian basin or
transitional forests. However, the material examined here was obtained from
a plant grown as ornamental on the Campus of the Universidade de Brasilia.
Several cercosporoid hyphomycetes are known from hosts of the Bombacaceae,
e.g. Cercospora bombacicola Munjal et al., C. bombacis Goh & W.H. Hsieh,
C. ceibae Chupp & Viégas, Pseudocercospora bombacina (T.S. Ramakr. & K.
Ramakr.) Deighton, P. eriodendri (Racib.) U. Braun (= Cercospora italica Curzi),
P. pachirae U. Braun et al. (= C. pachirae Chupp & A.S. Mull., nom. inval.),
Ramularia chorisiae Viégas (Chupp 1954, Vasudeva 1963, Braun 1998, Hsieh &
Goh 1990, Crous & Braun 2003). Silva & Minter (1995) listed 26 fungal species
on hosts of the family Bombacaceae studied by Batista and co-workers, among
them just 17 are hyphomycetes but none belonging in the Cercospora complex.
Three new cercoporoid species found on bombacaceous hosts have recently
been found in the Cerrado and are now described.
Cercospora ochromae A. Hern.-Gut. & Dianese, sp. nov. FIG. 4
MycoBank, MB 512038
Cercosporae ceibae similis, sed conidiis cylindraceis-obclavatis, basi obconice truncata.
SPECIMEN EXAMINED: BRAZIL. DISTRITO FEDERAL: BRASILIA, Campus of Universidade
de Brasilia, on living leaves of Ochroma sp. (Bombacaceae), 31 Jul 1994, leg. JC Dianese
1766, holotype (UB mycol. col. 6317).
Lesions 1-8 mm diam., amphigenous, small, punctiform, irregular, light
brown to brown, surrounded by a yellow halo. CoLontzs grayish, shiny, mainly
hypophyllous. Srromata poorly developed, superficial, textura globosa, 11-52
um diam. CoNIDIOPHORES densely fasciculate, straight, flexuous, 33-63
50 ... Hernandez-Gutiérrez & Dianese
Fig. 4. Cercospora ochromae on Ochroma sp. Lesions ona leaf (top center) among conidia,
around a dark stroma bearing typical Cercospora geniculate conidiophores with
conidiogenous cells showing darkened conidiogenous loci. Based on the holotype
(UB Mycol. Col. 6317).
um long, 4-6 um diam. at the widest portion, 0-2-septate, light brown, but
subhyaline at the apex, thin-walled, smooth. CONIDIOGENOUS CELLS 40-55 um
long, 2-3 um diam., integrated, polyblastic, sympodial, with 2-7 geniculations,
with thickened, darkened, prominent conidiogenous loci. Conip1A solitary,
straight, flexuous, sometimes curved, cylindrical to obclavate, obconically
Cercosporoids from the Brazilian Cerrado ... 51
truncate at the base with a thickened and darkened hilum, apex acute or round,
25-147 um long, 2-4 um diam. at the widest part, 2-3 um at the base and 1-2
uum near apex, 2—16-septate, subhyaline, smooth, thin-walled.
COMMENTS: Due to pigmented conidiophores, thickened and darkened
conidiogenous loci and solitary, hyaline conidia, Cercospora ochromae is a
typical Cercospora. It is close to C. ceibae, which is also characterized by having
rather short conidiophores with few septa, but the latter species differs from
C. ochromae in having acicular conidia with truncate base. On account of the
acicular conidia, Crous & Braun (2003) classified this species as part of the
Cercospora apii Fresen. complex. Cercospora bombacicola and C. bombacis are
two species with much longer, pluriseptate conidiophores and acicular conidia
with truncate base, reduced to synonymy with C. apii s. lat. by Crous & Braun
(2003).
Pseudocercospora pseudobombacis A. Hern.-Gut. & Dianese, sp. nov. FIG. 5
MycoBank, MB 512039
Differt a Pseudocercospora bombacina, P. eriodendri et P. pachirae conidiis longioribus
et imprimis distincte latioribus, 27-95 x 5-9 um, crassitunicatis, 1-14 transverse et 1-2
oblique septatis.
SPECIMEN EXAMINED: BRAZIL. DISTRITO FEDERAL: PLANALTINA, Esta¢gao Ecoldgica
de Aguas Emendadas, on living leaves of Pseudobombax sp. (Bombacaceae), 27 Feb 1997,
leg. M. Sanchez 2345, holotype (UB mycol. col. 13558).
LESIONS irregular, sometimes angular, amphigenous, brown, without a well-
defined margin, 1-14 mm diam., sometimes coalescent reaching up to 5.5
cm diam. COLONIES amphigenous, caespitose, dark brown or black, scattered
on lesions, erumpent or stomatal. MycELIUM immersed in the substrate.
STROMATA poorly developed. CONIDIOPHORES loosely fasciculate, lageniform
or ampulliform and without septa when young, cylindrical, lageniform and
1-5-septate when mature, straight or flexuous, smooth, geniculate, sometimes
sympodially proliferating, 35-55 um long, 2-4 um at the apex, 6-8 um diam.
close to the apex, 7-12 um at the base, light yellowish green to olivaceous.
CONIDIOGENOUS CELLS integrate, terminal, polyblastic, sympodial, geniculate;
conidiogenous loci inconspicuous, neither thickened nor pigmented. CONIDIA
solitary, straight, curved or flexuous, obclavate, truncate to obconically truncate
at the base, rounded at the apex, sometimes with a constriction near the base;
light green or yellowish green, paler at the apex, sometimes more intensely
pigmented, up to olivaceous, and thick-walled, except for the lighter basal and
apical cells, 27-95 um long, 5-9 um diam. at the widest part, 2-4 um at the base,
3-5 um at the apex, with 1-14 transverse and 1-2 oblique septa, smooth.
CoMMENTs: This fungus has some characteristics of Prathigada Subram.
species (thick and dark conidial walls and septa) but the conidiogenous loci
52 ... Hernandez-Gutiérrez & Dianese
5 um
40 pm
Fig. 5. Pseudocercospora pseudobombacis on Pseudobombax sp. Coalescent brown lesions
on a leaf (top left); conidia with thick walls and septa similar to those of Prathigada
species; sympodially proliferating septate conidiophores with conidiogenous cells
showing unthickened conidiogenous loci. Based on the holotype (UB Mycol. Col.
13558).
are inconspicuous as in Pseudocercospora species, so that this species has to be
placed in the latter genus. Several species of Pseudocercospora on hosts of the
Bombacaceae are known, but P. bombacina, P. eriodendri, and P. pachirae are
easily distinguishable from P pseudobombacis by their shorter and above all
much narrower, thin-walled conidia with few, always transverse septa (Chupp
1954, Vasudeva 1963).
Cercosporoids from the Brazilian Cerrado ... 53
Pseudocercospora eriothecae A. Hern.-Gut. & Dianese, sp. nov. FIG. 6
MycoBank, MB 512040
Differt a Pseudocercospora bombacina, P. eriodendri et P. pachirae conidiophoris aeque
brevibus et conidiis distincte septatis, rectis vel leviter curvatis.
SPECIMEN EXAMINED. BRAZIL. MINAS GERAIS: DIVINOPOLIS, Feira Agropecuaria, on
living leaves of Eriotheca sp., (Bombacaceae), 26 Jun 1995, leg. JC Dianese 2420, holotype
(UB mycol. col. 9079).
LesIons 1-14mmdiam.,amphigenous, irregular, coalescent, brown, surrounded
by a darker halo. CoLonies epiphyllous, grayish, scattered on the lesions.
MyceELIum immersed. STROMATA well-developed, subcuticular or immersed in
the epidermis, 12-42 um diam., mixed texture (textura globosa and angularis),
brown. CONIDIOPHORES in small, loose fascicles, shape variable, unbranched,
10-59 x 3-5 um, pale brown, thin-walled, smooth, 0-2-septate, sometimes
reduced to conidiogenous cells; comprising three types, depending on the state
of development: a) short conidiophores — lageniform, 10-17 um long, 3-5 um
diam. at the widest part, 1-2 um at the apex, aseptate (conidiophores reduced
to conidiogenous cells), sometimes sympodially proliferated; b) medium
conidiophores — lageniform to cylindrical, straight or slightly curved, pale
brown, 17-30 um long, 3-4 um diam. at the widest part, 1-2 um diam. at the
apex, 0-2-septate, with a single sympodial proliferation; c) long conidiophores
— cylindrical, straight, pale brown, smooth, flexuous or curved, 31-59 um long,
3-3.5 um diam. at the widest part, 1-3-septate, sympodial proliferations with
up to three geniculations per conidiophore. CONIDIOGENOUS CELLS integrated,
terminal, conidiogenous loci inconspicuous. ConipIA solitary, straight,
flexuous or curved, sometimes reaching almost 90° just below the second basal
septum, cylindrical or slightly obclavate, truncate at the base and rounded at
the apex, 27-63 um long, 2-3 um diam. at the widest part, 1-2 um near the
apex and at the base, sparsely and obscurely 0-6-septate, subhyaline, smooth,
thin-walled.
COMMENT: Pseudocercospora eriothecae is quite distinct from P. pseudobombacis,
described above, by its much narrower conidiophores and conidia with few,
consistently transverse septa. Pseudocercospora bombacina, P. eriodendri and P.
pachirae are similar, but possess uniformly short conidiophores and straight to
only somewhat curved conidia with conspicuous septa (Chupp 1954, Vasudeva
1963).
Cercosporoid fungi on Burseraceae
Twenty plant genera of tropical distribution belong in the family
Burseraceae, with three genera present in the Cerrado: Commiphora, Protium
and Tetragastris, with a total of 12 species, seven of them belonging in Protium
(Mendonga et al. 1998). Silva & Minter (1995) listed 109 fungal species on hosts
54 ... Hernandez-Gutiérrez & Dianese
Fig. 6. Pseudocercospora eriothecae on Eriotheca sp. Scattered black lesions on a leaf (top
center) among conidia with truncate, thin hila; erumpent stromata with textura
globosa, bearing conidiophores and conidiogenous cells with inconspicuous, flat
conidiogenous loci. Based on the holotype (UB Mycol. Col. 9079).
of the Burseraceae, among them 80 species occurring on species of Garuga
and Protium, including 24 hyphomycetes, but none of them belonging to the
Cercospora complex. Dianese et al. (1997) listed for the Cerrado nine species
of ascomycetes and four hyphomycetes, all of them occurring on Protium sp.
Few cercosporoid species have been described from hosts of this plant familiy,
Cercosporoids from the Brazilian Cerrado ... 55
viz. the Indian Cercospora boswelliae Harsh et al. on Boswellia serrata Roxb. ex
Colebr., C. burserae Govindu & Thirum. on Bursera delpechiana (DC.) Engl.,
and C. garuaicola Narayan et al. and Stenella garugae J. M. Yen et al. on Garuga
pinnata Roxb. (Yen et al. 1982, Crous & Braun 2003). The following new species
represents the first member of the genus Pseudocercospora on a host of the
Burseraceae.
Pseudocercospora protii A. Hern.-Gut. & Dianese, sp. nov. Fic. 7
MycoBank, MB 512041
Laesiones 5-10 mm diam., amphigenae, orbiculares, angulares vel irregulares, adaxiale
griseolae, margine brunneo-rubricoso cinctae, abaxiale brunneae, sine margine distincto.
Coloniae amphigenae, punctiformes. Stromata 60-140 um diam. ex texturis intermixtis
globosis et angularis, bene evoluta, subepidermalia, erumpentia, brunnea. Conidiophora
in fasciculis compactis aggregatis, recta, curvata vel flexuosa, cylindrica, 57-80 um longa,
4-5 um lata in parte latissima, 3—6-septata, brunneo-olivacea, apicem versus pallidiora.
Cellulae conidiogenae in conidiophoris incorporatae, terminales, praecipue monoblasticae,
aliquando polyblasticae, sympodiales; cicatricibus inconspicuis. Conidia solitaria,
filiformia vel anguste obclavata, curvata, subhyalina, guttulata, basi obconico-truncata,
hilo tenui claro, apice obtuso vel acuto, 55-201 um longa, 2-3 um lata in parte latissima,
1-2 um lata ad basem, 1-1.5 um lata prope apicem, 5—10-septata, laevia.
SPECIMEN EXAMINED: BRAZIL. GotAs: CrisTALina, Fazenda Nova India, on living
leaves of Protium ovatum (Burseraceae), 10 Apr 1993, leg. JC Dianese 831, holotype (UB
mycol. col. 3739).
LESIONS amphigenous, circular, angular or irregular, gray, with a reddish brown
halo on the adaxial surface, brown and without halo on the abaxial surface.
COLONIES amphigenous, punctiform, brown. STROMATA subepidermal, well-
developed, 60-140 um diam., mixed textura globosa and angularis, erumpent,
brown. CONIDIOPHORES densely fasciculate, numerous, straight, curved or
flexuous, cylindrical, 57-80 um long, 4-5 um diam. at the widest part, 3-6-
septate, brown-olivaceous, paler towards the apex, thin-walled, smooth.
CONIDIOGENOUS CELLS integrate, terminal, mainly monoblastic, occasionally
polyblastic, sympodial. COoNIDIOGENOUS LOCI inconspicuous, neither
thickened nor pigmented. Conrp1a solitary, filiform to narrowly obclavate,
curved, subhyaline, guttulate, obconically truncate at the base, obtuse to acute
at the apex, 55-201 um long, 2-3 um diam. at the widest part, 1-2 um at
the base, 1-1.5 um near the apex, 5-10-septate, smooth, thin-walled, hilum
unthickened, not darkened.
Cercosporoid fungi on Celastraceae
The family Celastraceae has about 60 genera worldwide, mainly in the tropics
and subtropics. The most frequent genera in the Cerrado are Maytenus with 12
species, Austroplenckia with only one species [A. populnea (Reissek) Lundell]
and Tontelea with 15 species (Mendonga et al. 1998). Numerous cercosporoid
56 ... Hernandez-Gutiérrez & Dianese
sensi tate see pa a cae
a |
pisepinese
Ce
sevethe
ran)
=
=
i
Fig. 7. Pseudocercospora protii on Protium ovatum. Brown coalescent lesions on a leaf
(top left); subepidermal erumpent stroma with textura angularis bearing septate
conidiophores and conidiogenous cells without conspicuous conidiogenous loci;
conidia pluriseptate pale brown. Based on the holotype (UB Mycol. Col. 3739).
hyphomycete species have been described on hosts belonging to the
Celastraceae, e.g. Cercospora celastricola Govindu & Thirum., C. euonymigena
Y.L. Guo & Y. Jiang, Cercosporella euonymi Erikss. (= Cercospora euonymi
Ellis), Mycocentrospora verrucosa Pollack & Ellett, Passalora melanochaeta
(Ellis & Everh.) U. Braun, Pseudocercospora celastri S. Singh, P. elaeodendri
Cercosporoids from the Brazilian Cerrado ... 57
(G.P. Agarwal & Hasija) Deighton, P destructiva (Ravenel) Y.L. Guo & X,J.
Liu, P gymnosporiae R.K. Dubey et al., P myricacearum M.K. Khan et al.,
P. paradoxa U. Braun & Bagyan., Pseudocercosporella gymnosporiae Bagyan. et
al... Ramularia celastri Ellis & G. Martin, Stenella celastri A.N. Rai & Kamal,
S. elaeodendri Kamal et al., S. hippocratiae K. Srivast. et al., Verrucisporota indica
(Kamal & P. Kumar) U. Braun (Chupp 1954, Deighton 1976, Singh 1979, Dubey
et al. 1990, Rai & Kamal 1990, Khan et al. 1991, Guo & Hsieh 1995, Braun 1995,
1998, Bagyanarayana & Braun 1999, Crous & Braun 2003). Dianese et al. (1997)
mentioned an ascomycetous species and a non-cercosporoid hyphomycete
occurring on Austroplenckia sp., while Silva & Minter (1995) listed 24
ascomycetous species and 19 non-cercosporoid hyphomycetes associated with
hosts of the family Celastraceae.
Two new cercosporoid hyphomycetes have recently been found on
A. populnea in the Cerrado, including one belonging in Pseudocercospora and
a second species which represents the first member of Prathigada on a host of
the Celastraceae.
Prathigada austroplenckiae A. Hern.-Gut. & Dianese, sp. nov. Fic. 8
MycoBank, MB 512042
Differt a omnibus speciebus generis conidiophoris ex cellulis stromaticis cylindraceis
oriundis et cicatricibus valde protuberantibus et fuscatis.
SPECIMEN EXAMINED: BRAZIL. DIsTRITO FEDERAL: BRASILIA, Centro Olimpico da
Universidade de Brasilia, on living leaves of Austroplenckia polpunea (Celastraceae), 26
Aug 1994, leg. C.A. Indcio 8, holotype (UB mycol. col. 17810); 11 May 1994, leg. C.A.
Inacio 114, paratype (UB mycol. col. 6109).
Lesions amphigenous, 1-3 mm diam., circular to elliptical, dispersed or
aggregated and coalescent to form lesions of up to 30 mm diam., dark brown.
COLONIES amphigenous, orange-brown, subcuticular or subepidermal,
erumpent. Stromata well-developed, textura globosa, 30-52 um diam.
CONIDIOPHORES arising from cylindrical stromatic cells, caespitose, straight,
flexuous or curved, lageniform, aseptate and witha single apical scar when young
or cylindrical, 1-4-septate, 23-40 um long, 4-7 um diam. in the widest part,
with enteroblastic rejuvenation (monopodial growth by rupture of the apical
region). CONIDIOGENOUS CELLS integrated, terminal, polyblastic, sympodial,
strongly cicatrized, with 1-30 darkened, thickened, strongly protuberant
conidiogenous loci. Conip1A solitary, simple, obclavate, sometimes rostrate,
truncate to obconically truncate at the base with a prominent, thickened,
darkened hilum, straight, flexuous, cylindrical to obclavate, 34-92 um long,
5-7 um diam. at the widest part, 2-3 um at the base, 2-4 um at the light
brown, rounded apex, 0-14-septate, septa and walls thick and dark, smooth or
sometimes finely verruculose.
58 ... Hernandez-Gutiérrez & Dianese
Fig. 8. Prathigada austroplenckiae on Austroplenckia populnea. Leaf with small lesions
that may coalesce (top left); conidia with thick, dark walls, septa, and hilum;
conidiophores sympodial, septate, with condiogenous cells strongly cicatrized
with dark conidiogenous loci; subcuticular erumpent stroma with textura
angularis. Based on the holotype (UB Mycol. Col. 17810).
Cercosporoids from the Brazilian Cerrado ... 59
COMMENT: This species has in common with the type species, Prathigada
crataevae (Syd.) Subram. (Subramanian & Ramakrishnan 1956), the presence
of subcuticular, well-developed stromata, sympodial conidiogenous cells and
sometimes rostrate conidia. However, the new species differs in possessing
conidiogenous loci more prominent than described in the type species, and
profusely geniculate conidiophores. These characteristics are also present in
P. terminaliae (Syd.) B. Sutton (Sutton 1994), but in this species the
conidiogenous loci are just slightly protuberant and the geniculations are less
distinct. Moderately protuberant loci are also present in P. maclurae (Thiim.)
U. Braun (Braun 1996). Geniculate conidiogenous cells and slightly thickened
scars also occur in P backmanii Furlan. & Dianese (Furlanetto & Dianese
1999). In general, conidia of all known Prathigada species are obclavate to
subcylindrical or fusiform, with some thick septa and thick cell walls. Three
species (P. crataevae, P. maclurae and P. terminaliae) have in common with the
new P. austroplenckiae the presence of slightly rostrate and verruculose conidia,
but stromata are absent or only small. However, the two main distinguishing
characters are the very dark protuberant scars on the conidiogenous cells, and
the presence of columns of almost cylindrical stromatic cells giving rise to the
conidiophores. These characters are not present in the species mentioned above,
as well as in all other previously known species, including P bauhiniae P. Rama
Rao, P. Rag. Rao & Manohar. (Rao & Manoharachary 1975), P. condensata
(Ellis & Kellerm.) U. Braun (Braun 1999), P gymnocladii (Ellis & Kellerm.)
U. Braun (Braun 1996), P. punjabensis (Subram.) Subram., P. tamarindi
Muthappa (Muthappa 1965) and in P zizyphi A.V. Rao & K. Ramakr. (Rao &
Ramakrishnan 1965).
Pseudocercospora austroplenckiae A. Hern.-Gut. & Dianese, sp. nov. FIG. 9
MycoBank, MB 512043
Laesiones parvae, 1-3 mm diam., amphigenae, orbiculatae, ellipticae, dispersae vel
aggregatae, coalescentes, usque ad 30 mm diam., atro-brunneae. Coloniae hypophyllae,
nigrae, nitidae, sparsae. Stromata 20-30 um diam., e texturis globosis, parva, rudimentalia,
substomatalia. Conidiophora in fasciculis laxis aggregata, brevia, unicellularia,
lageniformia, ampulliformia, laevia, olivacea, tenuitunicata, 10-17 um longa, 3-7 um
lata in parte latissima, 1-2 um lata prope apicem, vel conidiophora solitaria ex hyphis
superficialibus oriunda, cylindrica vel doliiformia-lageniformia, apice truncato, 6-13 um
longa. Cellulae conidiogenae integratae, terminales, monoblasticae, aliquando sympodiales;
cicatricibus inconspicuis, non incrassatis, non fuscatis. Conidia solitaria, simplicia,, recta,
curvata vel leviter flexuosa, cylindrica, obclavata, basi truncata vel obconico-truncata,
apice rotundato vel leviter acutato, 16-66 um longa, 2-4 um lata in parte latissima, 1-2
um lata ad basem et prope apicem, obscure 0-6-septata, olivacea, laevia, tenuitunicata.
SPECIMEN EXAMINED: BRAZIL. DisTRITO FEDERAL: BRAS{LIA, Centro Olimpico da
Universidade de Brasilia on living leaves of Austroplenckia polpunea (Celastraceae), 26
Aug 1994, leg. C.A. Indcio 8, holotype (UB mycol. col. 1189).
60 ... Hernandez-Gutiérrez & Dianese
Fig. 9. Pseudocercospora austroplenckiae on Austroplenckia populnea. Leat showing
brown irregular lesions; conidia obclavate, with thin septa and wall; fascicle of ,
conidiogenous cells arising from a substomatal stroma; superficial hypha with
conidiogenous cells. Based on the holotype (UB Mycol. Col. 1189).
Lesions mostly small, 1-3 mm diam., amphigenous, circular, elliptical, irregular,
dispersed or aggregated, confluent, reaching up to 10 mm diam., dark brown.
Cotonies hypophyllous, shiny black, sparse. STROMATA poorly developed,
Cercosporoids from the Brazilian Cerrado ... 61
20-30 um diam., textura globosa, substomatal. CONIDIOPHORES caespitose as
well as solitary, fasciculate conidiophores short, reduced to conidiogenous cells,
lageniform, ampulliform, emerging through stomata, olivaceous, smooth, thin-
walled, 10-17 um long, 3-7 um diam. in the widest part, 1-2 um near the apex;
solitary conidiophores laterally arising from superficial hyphae, cylindrical or
doliiform-lageniform, truncate at the apex, 6-13 um long. CONIDIOGENOUS
CELLS sometimes sympodially proliferating, conidiogenous loci inconspicuous.
Conip1A solitary, simple, straight, curved or slightly flexuous, cylindrical,
obclavate, truncate to obconically truncate at the base, rounded to slightly
acute at the apex, olivaceous, 16-66 um long, 2—4 um diam. at the widest part,
1-2 um at the base and near the apex, 0-6-septate, but septa rather obscure,
smooth, thin-walled, hila neither thickened nor darkened.
COMMENT: Several Pseudocercospora species are known on hosts of the
Celastraceae, e.g. P. celastri, P. destructiva, P. elaeodendri, P. gymnosporiae
and P myricacearum (Chupp 1954, Deighton 1976, Singh 1979, Dubey et al.
1990, Khan et al. 1991, Guo & Hsieh 1995), but all of them are characterized
by having well-developed stromata, much longer, septate conidiophores and
lacking superficial mycelium and solitary conidiophores. Pseudocercospora
austroplenckiae is distinguished from these species by its poorly developed
substomatal stromata and aseptate conidiophores, either fasciculate or solitary,
arising from superficial hyphae. Furthermore, P. celastri and P. gymnosporiae
possess much wider conidia, 2.5-10 um. The Indian P. paradoxa (Bagyanarayana
& Braun 1999) described from Elaeodendron sp., forms superficial hyphae, but
is quite distinct by having consistently solitary conidiophores, arising from the
superficial hyphae, up to 200 um long and 3-7 um wide, pluriseptate, and broad
conidia, 15-55 x 4-8 um, occasionally with 1-2 longitudinal or oblique septa.
Acknowledgments
The authors thank CNPq and Fundacao Banco do Brasil for financial support, and
Mariza Sanchez for assistance with the herbarium work. Thanks are also given to Uwe
Braun, whose extra help is much appreciated, and Roland Kirschner for pre-submisson
reviews that greatly contributed towards the improvement of this paper.
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Volume 106, pp. 65-67 October-December 2008
Checklists of the myxomycetes, larger ascomycetes,
and larger basidiomycetes in Turkey
ERTUGRUL SESLI* & CVETOMIR M. DENCHEV’
ertugrulsesli@ktu.edu.tr
‘Department of Biology Education, Karadeniz Technical University
Trabzon, Turkey
cmdenchev@yahoo.co.uk
"Institute of Botany, Bulgarian Academy of Sciences
23 Acad. G. Bonchev St., 1113 Sofia, Bulgaria
Abstract—This paper provides a compilation of available data on the myxomycetes,
larger ascomycetes, and larger basidiomycetes reported from or known to occur in
Turkey. Three main lists of correct names of myxomycetes, larger ascomycetes, and
larger basidiomycetes, recognized as occurring in Turkey, are given, in which the taxa
are alphabetically arranged. For each taxon, references are cited. An index of synonyms
based on literature records from Turkey is appended. A list of excluded records,
providing reasons for their exclusion, is also given. The complete checklists are available
on: http://www.mycotaxon.com/.
Key words—biodiversity, fungal diversity, macrofungi, Turkish mycota
Introduction
A number of studies on the mycota of Turkey (Fig. 1) have been published.
Contemporary knowledge on the diversity of Turkish myxomycetes and
macrofungi is based on a period of more than 90 years of investigations. In
the course of gathering literature sources, we found more than 400 scientific
works, which contain various Turkish records of myxomycetes and macrofungi.
During the 1990s and part of the first decade of 2000, the accumulation of a
huge number of records, without a published catalogue of the macrofungi in
Turkey, allowed room for many mistakes, including duplication of errors in
some articles concerning new Turkish records. In the past, some species were
published as new records many times and in different years by the same or
different authors, and in the same or different journals.
Author for correspondence
66 ... Sesli & Denchev
BULGARIA - Ss e a
o .
Edirne Sinep ©
: © Artvin
istanbul “S Samsun o ;
o Bolu Ord o 3 Trabzon ARMENIA ff
Marmara Sea ” ; Amasya rau fe)
o Koceah ps Giresun
o Gumiushane
O Sursa Ankara
Babkesir R he eee oErzurum ‘sd >
<4 °
ee Sait Lake Mus
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© Mugla o : Santiurfa .
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Fig. 1. Neighbouring countries and some of the major cities of Turkey
The first contribution to the macrofungi from Turkey was published by K.
Vlaev (1915). Sesli & Baydar (1995, 1996) presented the first checklists of
macrofungi, including Russulaceae and Agaricales, respectively. The first
checklist of myxomycetes was compiled by Ergiil & Dilger (2000). Sesli &
Denchev (2005) published a checklist of myxomycetes and the first complete
checklist of Turkish macrofungi and their synonyms. Additional checklists of
myxomycetes (Dilger 2007) and macrofungi (Solak et al. 2007) have recently
appeared.
The aim of this new edition of the checklists is a further attempt to summarize
and completely present the correct names of species of myxomycetes, larger
ascomycetes, and larger basidiomycetes currently known in Turkey, as well as
to list their synonymous names occurring in the available sources on the fungi
in Turkey. We hope that this paper will be a guide for future studies and a
helpful source for creation of a database of the Turkish mycota.
We intend to update regularly the Internet version of the checklists.
Materials and methods
Main lists of myxomycetes, larger ascomycetes, and larger basidiomycetes
have been developed. The taxa are given in alphabetical order. The numbers
within square brackets, following the authors of each species or infraspecific
taxon, refer to the reference(s) where the taxon is reported. A number [001-
288] and letters (e.g., a, b, c) are assigned to each reference in the section
“Bibliography of the contributions to myxomycetes, larger ascomycetes, and
Macrofungi and myxomycetes of Turkey ... 67
larger basidiomycetes in Turkey’. The generic and species treatment follows
many of the recent monographs and particular articles on the European fungi.
The names of authors of fungal taxa are abbreviated according to Kirk & Ansell
(1992) and Kirk et al. (2004).
Because many species have been published under different names, a thesaurus
of synonyms is separately listed with references to the correct names used in
the main lists. A list of excluded records, providing reasons for their exclusion,
is also appended.
Results
The complete checklists and estimation of the species, recognized as occurring
in Turkey, are available on: http://www.mycotaxon.com/. We envisage the
Internet version of the checklists to be regularly updated.
Acknowledgements
We gratefully acknowledge Dr Vladimir Antonin (Department of Botany, Moravian
Museum, Brno, Czech Republic), Prof. Timothy J. Baroni (Department of Biological
Sciences, College at Cortland, State University of New York, USA), Dr Roy E. Halling
(Institute of Systematic Botany, The New York Botanical Garden, Bronx, NY, USA), and
Prof. Steven L. Stephenson (University of Arkansas, Fayetteville, Arkansas, USA) for
critically reading the manuscript and serving as pre-submission reviewers. The work of
the senior author was supported in part by the Karadeniz Technical University (Project
mOn212116.001°2):
Literature cited
Dilger B. 2007. Checklist of the myxomycetes in Turkey. Mycologia Balcanica 4: 151-155.
Ergiil CC, Dilger B. 2000. Myxomycetes of Turkey. Karstenia 40: 39-41.
Kirk PM, Ansell AE. 1992. Authors of fungal names. International Mycological Institute, CABI,
Wallingford.
Kirk PM et al. 2004. Authors of fungal names. CABI Bioscience, Wallingford. Electronic version:
http://www.speciesfungorum.org/AuthorsOfFungalNames.htm.
Sesli E, Baydar S. 1995. A preliminary checklist of Russulaceae of Turkey. Russulales News 5: 5-22.
Sesli E, Baydar S. 1996. A preliminary checklist of Agaricales of Turkey. Mycotaxon 60: 213-224.
Sesli E, Denchev CM. 2005. Checklists of the myxomycetes and macromycetes in Turkey. Mycologia
Balcanica 2: 119-160.
Solak MH, Isiloglu M, Kalmis E, Alli H. 2007. Macrofungi of Turkey. Checklist. Vol. 1. Universiteliler
Ofset, Bornova-Izmir.
Vlaev K. 1915. [Contribution to the higher fungus flora of Turkish Thrace]. Travaux de la Société
Bulgare des Sciences Naturelles 8: 199-207. (In Bulgarian)
EEE EE EE EEE in
MY COTAXON
Volume 106. pp.69-74_ October-December 2008
Comatricha argentinae, a synonym of C. tenerrima
GABRIEL MORENO *, CARLOS ILLANA * & JORGE R. DESCHAMPS ”
gabriel.moreno@uah.es; carlos.illana@uah.es
' Dpto. de Biologia Vegetal, Facultad de Biologia, Universidad de Alcala
Alcala de Henares 28871, Madrid, Spain
? Universidad de Belgrano, Facultad de Ciencias Agrarias
Federico Lacroze 1955, Cap. Fed. 1426 Buenos Aires, Argentina
Abstract — The type material of Comatricha argentinae was studied. The type is the
only known collection of this species, which was described from Argentina. Sporocarp
morphology and the typical spore ornamentation under SEM coincide with those of
C. tenerrima. Thus, the synonymy of these species is proposed. Light microscope images
of sporocarps and capillitium along with scanning electron microscope photographs of
spore ornamentation are presented.
Key words — Myxomycetes, Stemonitales, taxonomy
Introduction
Comatricha argentinae is a species known only from its original description.
It was proposed as new to science by Deschamps (1974), based on a collection
from the decaying wood of Ulmus procera. In the original publication, the
species is described as Comatricha argentina. Later, Lado (2001) corrected the
name to Comatricha argentinae. According to the GBIF (www.gbif.net) data
base, no other records are known worldwide.
The first description included black and white drawings of the sporocarps,
which are similar to those of a number of other species of Comatricha. Line
drawings of the spores and capillitium were also included. Although the
capillitium is described as flat and provided with warts or rings, this appears
to be an unusual type of ornamentation for the genus. This is the reason why
we believed Comatricha argentinae needed detailed investigation by electron
microscopy.
Materials and Methods
The type material studied is located in the Herbarium of the Department of
Plant Biology, University of Alcala, Madrid, Spain (AH).
70 ...Moreno, Illana & Deschamps
The sporocarps studied were mounted in Hoyer’s medium. Spores were measured
to include surface structures such as spines or warts and measurements were
made with an oil immersion lens. For ultramicroscopic studies, the material to
be examined was rehydrated in concentrated ammonium hydroxide (28-30%)
for 30 minutes, dehydrated in aqueous ethanol (70%) for 30 minutes, fixed for
2 hours in pure ethylene glycol dimethyl ether (= 1,2-dimethoxymethane)
and finally immersed in pure acetone for at least 2 hours. This was followed by
critical point drying and sputtering with gold-palladium.
The micrographs were taken at the University of Alcala using a Zeiss DSM-950
microscope. The technique followed requires the use of very little material (one
sporocarp, a part of it or only a small portion of spores). Terminology used
herein to describe spore ornamentation follows that of Rammeloo (1975a, b).
Description
Comatricha argentinae J.R. Deschamps, Bol. Soc. Argent. Bot. 15(4): 340 (1974)
EiGsal 7
Fructificationes solitariae vel gregariae ex hypotalum minutum badium fulgidae
conspiquum surgentes. Esporamgia castanea, elliptico-ovoidea vel globosa, superne
attenuata, 0.3-0.6 x 0.5-1.2 mm, peridio obscuriore, plerunque evanescente sed in base
esporangii persistente et calyculo circulare castaneo-rufescente simulante. Pes nigrescens
vel badius opacus, subulatus, cavus, longitudinaliter substriatus quam espongium 2-3
vices longior. Columnella plerunque exserta, % partibus longitudinis vel toto sporangio
occupandum, percurrens badia, apice leviter ramosa et in filiis tenuibus desinentia
capillicium formans. Capillitium ex columnella vel raro ex calyculo surgente evanescente
plerunque ramosum cavum 1-4 um diam., laeve vel cum verrucis annulisve ornatum,
pallide castaneum. Esporarum massae pallide badiae, globosae, verrucosae, conspicuae
(6-)6.6-7.7(-8) um diam, Plasmodium ignotum.
HoLotypus — Argentina, Buenos Aires, Llavallol, Instituto Fitotécnico de Santa
Catalina, leg. L. Frias & G. Rovetta, 5-X-1970 (BAFC 22868 holotype), (LPS 37739
isotype), (BPI isotype).
HABITAT — On decaying Ulmus procera wood, after frequent rain.
EXAMINED MATERIAL — Material type BAFC 22868 is stored as AH 31762, along with
a Hoyer’s medium preparation.
The type material is stored in a match box with three pieces of wood of various
sizes glued to its the base. A card which has only three stalks and two columellae
and no capillitium or spores is stuck to the bottom of the box. The specimen
is very sparse, consisting of few complete sporocarps, some of which are very
hardened (Fig. 3), and only the stalks of some are preserved. However, a single
mature sporocarp (Fig. 4) was available for study of its morphology and spore
ornamentation under MO and SEM (Figs. 6-7).
Due to the impossibility of compiling a complete macroscopic description of this
species, the detailed description published by Deschamps (1974) is translated
Comatricha argentinae & C. tenerrima ... 71
2 d
N,°., DAF. 2266 3) ie Ode “Noosa ase Ax tae 3
UNIVERSIDAD DE BUENOS AIRES
4 FACULTAD DE CIENCIAS EXACTAS Y¥ NATURALES
DEPARTAMENTO DE CIENCIAS BIOLOGICAS
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cas Lew Bake cates, Ay — ty dba Qo- 08
i He Clan? Opal ;
ey
Boe Hida evox columate
at Sedu te
fran seta Coit age hades
4
Fics. 1-7 Comatricha argentinae (holotype BAFC 2 22868): 1. Original label. 2. Drawings kept
with the type sample. 3. Hardened sporocarp. 4. Mature sporocarp. 5. Spore. 6. Detail of spore
ornamentation. SCALE BARS. 3-4 = 1 mm. 5-6 = | pm.
72. ...Moreno, Ilana & Deschamps
as follows: “Solitary fructifications or in gregarious groups, appearing from a
small hypothallus, dark brown reddish, shiny, conspicuous. Brown sporangia,
elliptical-ovoid to spherical with the superior region (apex) slightly thinner due
to the inner presence of the columella, which frequently protrudes from the
tip of the sporotheca, 0.3-0.6 x 0.5-1.2 mm with a darker peridium, usually
evanescent, but persisting at the base of the sporotheca as a circular calyculus,
brown-reddish by transparency. Blackish to dark brown stalk, opaque, subulate,
hollow, somewhat striated lengthways, 2-3 times the length of the sporotheca.
Columella occupying all the sporotheca or up to % of its length, percurrent,
dark reddish brown, branching slightly at the extremities and producing thin
threads which form the capillitium. Capillitium arising from the columella and,
in some cases, the calyculus, evanescent, frequently branched, hollow, 1-4 um
diameter, flat or ornamented with warts or rings, light brown. Spores brown in
mass, light brown reddish under transmitted light, spherical, with conspicuous
groups of warts, (6—)6.6-7.7(-8) um. Unknown plasmodium.”
Discussion
Deschamps (1974) stated that Comatricha argentinae is similar to Collaria
rubens (Lister) Nann.-Bremek. (= Comatricha rubens Lister), indicating that
“this species has got characteristics which are similar to Comatricha rubens
because of its calyculus and the way the capillitium appears. However, the
sporangium’s colour, some capillitium details and the spores may indicate
that this is a new taxon for science”. Deschamps stated that the validity of the
new species had been confirmed by M.L. Farr, information that is cited in her
monograph (Farr 1976) but without dealing directly with the species, due to the
proximity in the time of publication of both investigations.
In a later work, Deschamps (1975) presented a key to the species of Comatricha
found in Argentina. In this key, he separated C. rubens and C. argentinae based
on sporocarp colour (more reddish in C. rubens) and spore ornamentation
(slightly spiny in C. rubens and with groups of prominent warts in C.
argentinae). Furthermore, he (Deschamps 1974) had previously indicated that
“the columella branching and the capillitium structure place Comatricha
argentinae near Lamproderma.”
In our revision of the type, we certainly place the taxon within the genus
Comatricha because of the sporocarps having a stalk with a fibrous base, a
columella that reaches the apex of the sporotheca and the winding branches of
the capillitium (Fig. 5), which appear all along the columella and are a somewhat
branched. Deschamps (1974) had noted that his species was closely related to
Lamproderma but evidence for this must inevitably now be refuted.
Comatricha argentinae & C. tenerrima ... 73
In our macro-photographs of Comatricha argentinae together with those taken
under SEM, it can be clearly seen that all the characteristics coincide with
Comatricha tenerrima, for the following reasons:
1. Comatricha argentinae and C. tenerrima produce stipitate sporocarps,
with a fusiform or ovoid light brown sporotheca.
2. In Comatricha argentinae and C. tenerrima the columella reaches the
apex of the sporocarp and gradually tapers upward. The capillitium is
abundant in both species, the branches are all the same in diameter,
flexuous, with a flat ornamentation and arise from the whole length of
the columella.
3. Under the optical microscope, both species share the same type of
spore ornamentation; the spores are warted, with more prominent
groups of warts. Under MEB, these warts resemble small baculi with
the apex divided into lobes and appearing star-like.
Rammeloo (1981) was the first to study the spore ornamentation of the
Comatricha tenerrima under MEB, and the latter was described as “trés
caractéristique pour cette espeéce’. Later, this ornamentation was confirmed on
many occasions by Moreno et al. (1993) and Lizarraga et al. (1999).
Comatricha tenerrimais notthe only taxon that has a special spore ornamentation
consisting of small baculi with stellate or lobed apices. Comatricha tenerrima
var. macrospora Rammeloo, a taxon known only from the Rwanda Mountains,
has a similar spore ornamentation but its sporocarps have longer stalks than
those of C. tenerrima, possess a peridial collar that subtends the sporotheca
and larger spores (Rammeloo 1981, Singer et al. 2006). Comatricha brachypus
(Meyl.) Meyl., is a similar species with a spore ornamentation consisting of
small baculi that fuse to form a pyramid (Singer et al. 2006).
Conclusion
According to the data presented above, we conclude that Comatricha argentinae
and C. tenerrima are two species with identical macro- and microscopic
characteristics; therefore, their synonymy is proposed:
Comatricha tenerrima (M.A. Curtis) G. Lister, Guide Brit. Mycetozoa, ed. 4: 39
(1919)
= C. argentinae J.R. Deschamps, Bol. Soc. Argent. Bot. 15(4): 340 (1974)
Acknowledgements
We wish to express our gratitude to Dr. S.L. Stephenson and Mr. D.W. Mitchell for
reviewing the manuscript and their useful comments and our thanks to Mr. A. Priego
and Mr. J.A. Pérez of the Electron Microscopy Service of the University of Alcala de
74 ...Moreno, Illana & Deschamps
Henares for their invaluable help with the SEM. We also thank Luis Monje of the
“Gabinete de Dibujo y Fotografia Cientifica” at the Universidad de Alcala de Henares
for his help in the digital preparation of the photographs, and we are grateful to Dr.
J. Rejos, curator of the AH herbarium.
Literature cited
Farr ML. (1976). Flora Neotropica. Monograph n°. 16. Myxomycetes. The New York Botanical
Garden, New York. 304 pp.
Lado C. (2001). Cuadernos de trabajo de Flora Micoldgica Ibérica 16. Consejo Superior de
Investigaciones Cientificas, Real Jardin Botanico de Madrid. 219 pp.
Lizarraga M, Illana C, Moreno G. (1999). SEM studies of the Myxomycetes from the Peninsula of
Baja California (Mexico), I. Arcyria to Fuligo. Ann. Bot. Fennici 35: 287-306.
Moreno G, Illana C, Burguete A. (1993). Spanish Myxomycetes. VU. (Province of Castellon).
Mycotaxon 46: 407-424.
Deschamps JR. (1974). Una nueva especie de Comatricha (Stemonitaceae-Myxomycetidae). Bol.
Soc. Argent. Bot. 15(4): 340-342.
Deschamps JR. (1975). Los Myxomycetes de la Argentina. Catalogo critico, distribucion y clave de
las especies. Physis, C. 34(89): 159-178.
Rammeloo J. (1975a). Structure of the epispore in the Stemonitales (Myxomycetes) as seen with the
scanning electron microscope. Bull. Jard. Bot. Belg. 45: 301-306.
Rammeloo J. (1975b). Structure of the epispore in the Trichiaceae (Myxomycetes) as seen with the
scanning electron microscope. Bull. Soc. Roy. Bot. Belgique. 107: 353-359.
Rammeloo J. (1981). Echinosteliales et Stemonitales (Myxomycetes). In: Flore illustrée des
champignons d Afrique Centrale. Fascicule 11: 214-244. Ministére de l’Agriculture. Jardin
Botanique National de Belgique.
Singer H, Moreno G, Illana C. (2006). SEM studies on type specimens of two rare Comatricha taxa.
Sydowia 58(2): 300-308.
MY COTA XON
Volume 106, pp. 75-102 October-December 2008
Additions to the myxomycete biota of Cuba. 1
M. CAMINO ', G. MORENO ”, A. CASTILLO 7,
D.W. MITCHELL 3 & D.W. MINTER *
hajb@ceniai.inf.cu
! Jardin Botanico Nacional de Cuba, Universidad de la Habana,
Carretera El Rocio, Km 3%, Calabazar, Boyeros,C.P. 19230, Ciudad Habana, Cuba
gabriel. moreno@uah.es, aurelio.castillo@uah.es
* Dpto. de Biologia Vegetal (Botanica), Universidad de Alcala
E-28871 Alcala de Henares, Madrid, Spain
slimelord@talktalk.net
>Walton Cottage, Upper Hartfield, East Sussex, England TN7 4AN
d.minter@cabi.org
*CABI, Bakeham Lane, Egham, Surrey, TW20 9TY, United Kingdom
Abstract — A study was carried out on the taxonomy and chorology of myxomycetes
based on field collections from various localities in Cuba. Some additional collections
were obtained from moist chamber cultures. Species recorded for the first time from
Cuba were Calomyxa metallica, Comatricha anomala, Craterium paraguayense,
Cribraria confusa, C. macrocarpa, Dictydiaethalium plumbeum, Diderma testaceum,
Didymium bahiense, D. difforme, D. nigripes, Licea pedicellata, L. pseudoconica,
Macbrideola cornea, Physarum decipiens, P. flavicomum, P. globuliferum, P. gyrosum,
P. mutabile, P. nicaraguense, P. oblatum, P. rigidum, P. roseum, P. stellatum, Reticularia
jurana and Trichia agaves. Ten species (Comatricha anomala, Craterium paraguayense,
Cribraria macrocarpa, Didymium bahiense, L. pseudoconica, Macbrideola cornea,
Physarum decipiens, P. gyrosum, P. mutabile and Trichia agaves) were new records for
the insular Caribbean. Species little studied by SEM or of particular interest for the
island’s biota are illustrated with SEM micrographs of spore ornamentation prepared
by critical point drying.
Key words — biodiversity, Myxomycota, Neotropical myxobiota
Introduction
The recent increase in biodiversity studies of myxomycetes from the tropics
and subtropics has taken place primarily in Central and South American
countries such as Costa Rica (Schnittler & Stephenson 2000), Ecuador
(Schnittler & Stephenson 2002, Stephenson et al. 2004) and Mexico (Lizarraga
76 ... Camino & al.
et al. 1997, 1998, 1999a,b, 2003, 2005a,b; Lado et al. 1999, 2003; Moreno et
al. 2006, Stephenson et al. 2003). For the insular Caribbean there has been
some exploration of Puerto Rico (Novozhilov et al. 2001), although the most
important recent study of myxomycete biogeography for the region is a general
compilation of fungi which also includes myxomycetes (Minter et al. 2001,
2002).
There are still, however, very few works relating to this group of organisms
from the Caribbean, despite an ample variety of ecosystems with a great diversity
of conditions and suitable substrates for their development. The present study,
using field collections and specimens obtained from moist chamber cultures,
was carried out because there is still very little published information about
myxomycetes in Cuba, the largest island of the region. Some species already
recorded from Cuba are also included where there are new locality records.
Materials and methods
Voucher material used in this study has been deposited in the myxomycete reference
collection of the Jardin Botanico Nacional, Habana (HAJB) and/or in some cases the ~
reference collection in Alcala de Henares (AH), and are from collections carried out
in different areas of the country. Specimens of myxomycetes fruiting under natural
conditions in the field were reviewed. The moist chamber culture technique (Stephenson
& Stempen, 1994) was used to reveal species associated with certain microhabitats,
especially the bark surfaces of living trees.
Samples for light microscopy were mounted in Hoyer’s medium or PVA following
Schnittler & Novozhilov (1996) and Koske & Tessier (1983). Spore measurements were
made using an oil immersion objective and include such surface structures as spines or
warts.
Scanning electron microscopy (SEM) images were prepared using critical point
drying (Castillo et al. 1997); the photomicrographs were produced at the University of
Alcala using a Zeiss DSM-950. This technique allows the use of very little material (part
of single sporocarp or even no more than a few spores).
The terminology used for the spore-producing stages follows Doérfelt & Marx (1990)
and Lado & Pando (1997). Spore wall ornamentation as seen by SEM is described
according to the terminology proposed by Rammeloo (1975a,b) and abbreviations
for author citations follow Kirk & Ansell (1992). The nomenclature used follows Lado
(2001) and Lado et al. (2005).
Species in the annotated list below are arranged alphabetically and new records for
Cuba are preceded by an asterisk. Records based on specimens harvested from moist
chamber cultures are indicated in each instance by “bc”. Numbers relating to material
supplied by David Mitchell are indicated by “DWM”.
The material considered in this study came from 58 localities. These localities are
listed under provinces and municipalities in order from west to east, with an identifying
number assigned to each (Map, TABLE 1).
Myxomycete biota of Cuba (1) ... 77
MapP 1. Localities of the material studied.
TABLE 1. Province, municipality, locality and species name of the material studied.
PROVINCE MUNICIPALITY
1-7.
PINAR .
; 1. VINALES
DEL Rio
4. LA PALMA
7. CANDELARIA
LOCALITY
1.1. km 17, Vifiales—Pons highway, to the right of the
4.2. Area Protegida de Recursos Manejados “Mil
Cumbres’, Reserva San Marcos
7.1. Reserva de la Biosfera “Sierra del Rosario, El Salén,
near the Academy house
8-9.
CIUDAD
DE LA HABANA
8.1. ARROYO NARANJO
8.2. BOYEROS
9g. PLAZA DE LA
8.1.1. Near the bridge of Calle 100 and Bejucal street
8.2.3. Jardin Botanico Nacional, “El Bambw’, near the
restaurant, 100 msm
9.1. Facultad de Biologia, yard
ee ee Oe REV OLUCION eek ae nee eee a ES 2 a ee ee
78 ... Camino & al.
TABLE 1, continued
LOLA ELAGAN TT on ROee 10.1. Approximately 2.3 km from the restaurant “El
Arabe”
ae 11. SANCTI SPIRITUS 11.1. Reserva Ecolégica “Alturas de Banao”, junction of
SANCTI SPfRITUS the Banao river, M* Antonia'’s farm
the Higuanojo river, the north of La Sabina biological
Pesta. allenic pie g1ove skeen eee ee ee ee
11.3. Reserva Ecoldgica “Alturas de Banao’, La Sabina
11.10. Reserva Ecolégica “Alturas de Banao’, La Sabina,
border of field station area
11.12. Reserva Ecoldgica “Alturas de Banao’, Las Damas,
ca dadianix’s farm) Giclee O08 tae Ce eee
11.13. Reserva Ecoldgica “Alturas de Banao’, Los Leones
stream, upper slopes between La Sabina and the ridge
11.14. Reserva Ecoldgica “Alturas de Banao’, on path
from M® Antonias farm to Ruinas de Boleto
11.16. Reserva Ecoldgica “Alturas de Banao’, stream at
the southwest base of Tetas de Juana, 600 msm
11.18. Reserva Ecoldégica “Alturas de Banao’, Tetas de
Juana, southwest hillside, 600 msm
11.20. Reserva Ecoldgica “Alturas de Banao’, route from
La Sabina biological station to El Regalo, spring area to
the north, 500 msm
11.22. Reserva Ecolégica “Alturas de Banao’, route from
La Sabina biological station to Hoyo del Naranjal, 2 km
from La Sabina biological station, 500 msm
12, CAMAGUEY 12. SIERRA DE CUBITAS 12.1 Hoyo de Bonet J
18-19. HOLGUIN
18. Mayari 18.1. La Mensura hill, base of south slope
19 Moa 19.1. Cayoguan River, left bank, near the campsite
14-15.GRANMA — 14. BUEY ARRIBA 14.1. Path to Pata de la Mesa, on the left toward the La
15. GUISA 15.1. Jardin Botanico Cupaynicut, bank of a river
Myxomycete biota of Cuba (1) ... 79
TABLE 1, concluded
13, 16-17. 13. GUAMA 13.1. Parque Nacional Turquino, path between Pico Cuba
SANTIAGO DE Peonciias Gucvas Seater seat ee eh EO eas
13.2. Parque Nacional Turquino, path between Pico Cuba
tuBs and Paso del Cadete, 1400 msm
13.3. Parque Nacional Turquino, path from Pico Cuba to
Las Cuevas, near Alto del Caldero, 1265 msm
16. GUAMA 16.1. Arroyon River, on the ascent of El Gato hill
17. SANTIAGO DE CUBA 17.1. La Isabelica
20.
Pur NA TiO 20. BARACOA 20.1. Path to Yunque de Baracoa
Annotated species list
* Calomyxa metallica (Berk.) Nieuwl., American Midland Naturalist 4: 335 (1916).
MATERIAL EXAMINED. LOCALITY 8.2.7, moist chamber culture on bark of Antidesma
montanum, 12-03-2003, leg. M. Camino, obtained by D.W. Mitchell, DWM 6810bc.
DisTRIBUTION: This is a new record for Cuba. In the insular Caribbean, the species was
known previously only from Jamaica (Minter et al. 2001).
* Clastoderma debaryanum var. debaryanum A. Blytt, Bot. Zeitung (Berlin)
38: 343 (1880).
MATERIAL EXAMINED. LOCALITY 13.5, on wood debris, 19—XI-2003, leg. M. Camino,
Le Ortiz & J. M Perez, HAJB'9795.
DISTRIBUTION: Together with samples from the Ecological Reserve “Alturas de Banao”
(Camino et al. 2007), this is the first record for Cuba. In the insular Caribbean this variety
has been recorded from Dominica, Guadeloupe and Jamaica (Minter et al. 2002).
COMMENTS - The sample is abundant and shows the characteristics of the
typical variety, well illustrated in Emoto (1977).
* Comatricha anomala Rammeloo, Bull. Jard. Bot. Natl. Belg. 46: 237 (1976).
MATERIAL EXAMINED. LOCALITY 8.2.7, moist chamber culture on bark of Jacaranda
arborea, 12-I1-2003, leg. M. Camino obtained by D.W. Mitchell, DWM 6840bc, HAJB
(dried specimens), AH (slide only).
DISTRIBUTION: This is a new record for Cuba and insular Caribbean.
COMMENTS — Comatricha anomala differs from other species of the genus in
its unique spore ornamentation. This taxon has been reported from several
localities in Europe and from California (USA), an unspecified location in
North America (Kowalski & Demaree 1987) and Baja California, Mexico
(Lizarraga et al. 1997).
80 ... Camino & al.
* Comatricha tenerrima (M.A. Curtis) G. Lister, Guide to Brit. Mycetozoa, ed. 4: 39
f19T9).
MATERIAL EXAMINED. LOCALITY 11.9, on rotten trunk of Samanea saman, 14-XI-2003,
leg. M. Camino & J. L. Ortiz, HAJB 9631.
DisTRIBUTION: In the insular Caribbean, this species has been recorded previously from
Jamaica (Minter et al. 2002).
COMMENTS - This species is characterized by an ovoid sporotheca, a pale
brown colour, a sharp apex to the sporotheca and a sinuous capillitium. This
collection, along with another three, also from the Ecological Reserve “Alturas
de Banao” (Camino et al. 2007) are the first records from Cuba.
Craterium leucocephalum var. scyphoides (Cooke & Balf. f.) G. Lister, in Lister,
Monogr. Mycetozoa, ed. 2: 97 (1911).
MATERIAL EXAMINED. LOCALITY 11.18, on dry twig, 26-I-1997, leg. M. Camino, HAJB
7632. LOCALITY 11.11, on dry leaf and twig, 18-09-1997, leg. U. Eliasson, M. Camino,
E. Bécquer & J. Calvino, HAJB 7809. Locaity 13.3, on wet leaf, 21-XI-2003, leg. M.
Camino, J. L. Ortiz & J. M. Pérez, HAJB 9843.
DIsTRIBUTION: C. leucocephalum s.]. has been recorded from Cuba, Dominica, Jamaica,
Puerto Rico and Trinidad & Tobago (Minter et al. 2002).
COMMENTS — Craterium leucocephalum var. scyphoides differs from C.
leucocephalum var. leucocephalum (Pers. ex J.F. Gmel.) Ditmar in having smaller
sporocarps and globose to subglobose sporothecae, where var. leucocephalum
has cylindrical or turbinate sporothecae. The spore ornamentation is also more
pronounced.
* Craterium paraguayense (Speg.) G. Lister,
in Lister, Monogr. Mycetozoa, ed. 2: 95 (1911). Fics. 1-3
MATERIAL EXAMINED. LOCALITY 13.3, on leaf and dry twig, 21-XI-2003, leg. M.
Camino, J.L. Ortiz & J.M. Pérez, HAJB 9844.
DISTRIBUTION: Craterium paraguayense was described originally from Paraguay (hence
its name), and was considered by Martin & Alexopoulos (1969) “to be largely subtropical
or tropical”. This is a new record for Cuba and the insular Caribbean.
COMMENTS ~ This species is characterized by stipitate sporocarps, cylindrical
violet sporothecae with apical circumscissile dehiscence, and cylindrical and
central pseudocolumellae with large calcium carbonate nodes of more or less
intense violet colour.
Castillo et al. (2002) made comparative studies of the types of Craterium
paraguayense and Physarum newtonii T. Macbr., for both macroscopic and
microscopic characters. The spore ornamentation, under both OM and SEM
show marked differences between the species. In C. paraguayense the spores
have small warts when viewed with the OM and short and narrow baculae when
viewed with the SEM; in P. newtonii there are very pronounced warts (OM) and
Myxomycete biota of Cuba (1)... 81
thick, prominent baculae (SEM). These characters were also observed in the
Cuban collection.
The Cuban specimen has violet to white-violet tones that look like discoloured
forms of Craterium leucocephalum var. scyphoides, but the black stipe and spore
ornamentation under SEM show the typical decoration of warts or short and
narrow baculae of C. paraguayense (Castillo et al. 2002).
* Cribraria confusa Nann.-Bremek. & Y. Yamam., Proc. K. Ned. Akad. Wet., Ser. C,
Biol. Med. Sci. 86(2): 212 (1983).
MATERIAL EXAMINED. LOCALITY 13.4, in moist chamber culture on bark of Cyrilla sp.,
leg. M. Camino, 20-XI-2003, obtained by D.W. Mitchell 2-03-2006, DWM 7048bc.
DISTRIBUTION: This is a new record for Cuba. In the insular Caribbean it has been
recorded previously from Dominica (Minter et al. 2002).
CoMMENTS - This species has the following characters: sporocarps small,
long-stalked, yellow and 0.3-0.7 mm in height; sporothecae small, globose,
0.05-0.2 mm diam., with a globose to subglobose; evanescent peridium, from
which threads originate forming a net of polygonal meshes up to 40 um diam.,
with narrow more or less isodiametric threads about 1 um diam., and slightly
flattened nodes with small and sparse mounds of calcium oxalate globules
present at the union of the threads; stalk hyaline to straw-yellow, 0.2-0.5
mm long, with granular material from half the height upwards; hypothallus
inconspicuous; spores are 5-7 um diam., globose to subglobose, warted and
with small crests, light yellow under OM.
Cribraria confusa was described by Nannenga-Bremekamp & Yamamoto
(1983), separating it clearly from C. minutissima Schwein. The two species
were compared using SEM by Keller et al. (1988). It was reported recently from
Chihuahua (Mexico) by Lizarraga et al. (2003).
Cribraria intricata Schrad., Nov. Gen. Pl.: 7 (1797).
MATERIAL EXAMINED. LOCALITY 12.1, on decayed wood with mosses, 16-05-95, leg. M.
Camino & R. Rankin, HAJB 7355. Locatity 7.1, on a very wet dead trunk, 07—X-1999,
leg. M. Benitez, HAJB 8568, DWM 6723bc (slide). Loca.iry 11.6, wet evergreen forest,
on decayed twig, 24—XI-2002, leg. M. Camino, HAJB 9415.
DIsTRIBUTION: This species is listed for Cuba by Farr (1976) from a specimen deposited
in FH; this was later confirmed by Camino et al. (2005).In the insular Caribbean, it
has been recorded from Cuba, Dominica, Jamaica, Puerto Rico and Trinidad & Tobago
(Minter et al. 2002).
ComMENTs - This species has a long stipe, 4/5 of the total sporocarp height.
The sporotheca has a cup with a calyculus with prominent, dark pulvinate
nodes filled with dictydine granules, each with radial, smooth and hyaline free
ends, 1-1.5 um diam. Spores are 5-6 um diam., ochraceous in mass, pallid
under OM.
82 ... Camino & al.
Fics. 1-3 Craterium paraguayense HAJB 9844, 1-2. Spores (bar = 2 um). 3. Detail of spore
ornamentation (bar = 1 ttm). Fics. 4-6. Didymium bahiense, HAJB 9827. 4. Detail of the lenticular
columella (bar = 50 um). 5. Spores (bar = 2 um). 6. Detail of spore ornamentation (bar = 1 um).
Cribraria tenella Schrad. is very close to C. intricata and is often difficult to
differentiate because of the presence of intermediate forms (Farr 1976). It
is frequent in North America and was cited from Cuba in 1869. It would be
interesting to investigate the possible synonymy of these two species and study
American and European collections.
Cribraria languescens Rex and C. stellifera Nowotny & H. Neubert are related
to Cribraria intricata. The first has coppery coloured sporothecae, flat nodes
and spores of 6.5—7.5 um; the second has larger spores of 9-10 um diam., and
is a temperate species.
Myxomycete biota of Cuba (1) ... 83
* Cribraria macrocarpa Schrad., Nov. Gen. Pl.: 8 (1797).
MATERIAL EXAMINED. LOCALITY 18.1, on bark of Pinus cubensis, 01-07-2004, leg. M.
Camino, HAJB 10322
DIsTRIBUTION: This is a new record for Cuba and the insular Caribbean. For the
Neotropics it has been reported previously only from Colombia and the USA (Florida)
(Farr 1976).
ComMENTSs - This species is characterized by its 0.5-0.8 mm diam. curved to
nodding olive-coloured sporothecae, and a 1-2 mm long black stipe. Empty
sporothecae have a pronounced transparency, with hollows and lines or smooth
approximately fusiform ribs; the upper part of the sporothecae have smooth
fusiform nodes, with abundant radial free ends. The 5-6 um diam. spores are
almost smooth and hyaline.
Cribraria microcarpa (Schrad.) Pers., Syn. Meth. Fung.: 190 (1801).
MATERIAL EXAMINED. LOCALITY 11.16, on bark of rotten trunk, 20-IX-97, leg. U.
Eliasson, M. Camino, E. Bécquer & J. Calvifio, HAJB 7895. Locatity 1.2, on rotten
trunk, 15-XII-2003, leg. M. Cabarroi, M. Camino & J. M. Pérez, HAJB 9917.
DisTRIBUTION: Together with the collections of Camino et al. (2007) from the Ecological
Reserve “Alturas de Banao’, these are the first records from Cuba. In the insular
Caribbean, this species has been recorded from Dominica, Guadeloupe, Jamaica, Puerto
Rico and Trinidad & Tobago (Minter et al. 2002).
CoMMENTS - This species is characterized by its small size, long and thin stipe,
peridial network with dark nodes and a small dark calyculus. This material was
gathered in the field from very wet substrata.
* Dictydiaethalium plumbeum (Schumach.) Rostaf. ex Lister, Monogr. Mycetozoa:
157 (1894).
MATERIAL EXAMINED. LOCALITY 11.4, on decayed wood, 28-I-2004, leg. M. Camino,
M. Cabarroi, A. Fidalgo, J.M. Pérez & T. Rojas, HAJB 10048. Locatiry 5.1, under bark
of very hard dry trunk, 19-III-2006, leg. M. Camino, HAJB 10863.
DISTRIBUTION: These specimens constitute the first records for Cuba. Previously in the
insular Caribbean, this species has been recorded from the American Virgin Islands, the
Dominican Republic and Puerto Rico (Minter et al. 2002).
ComMMENTSs - Dictydiaethalium plumbeum forms a depressed pseudoaethalium
consisting of closely compacted sporocarps. The upper peridium is tesselate. The
colour of the fructifications studied was yellowish brown; the other characters
were typical for this species.
Diderma effusum (Schwein.) Morgan, J. Cincinnati Soc. Nat. Hist. 16: 155 (1894).
= D. cubense Berk. & M.A. Curtis, in Berkeley, J. Linn. Soc. Bot. 10: 347 (1868).
MATERIAL EXAMINED. LOCALITY 13.1, on dry leaf, 20—XI-2003, leg. M. Camino
J. L. Ortiz & J.M. Pérez, HAJB 9829. Idem, HAJB 9830. Locauity 11.14, on dry
leaf, 30-I-2004, leg. M. Camino, M. Cabarroi, A. Fidalgo, J.M. Pérez & T. Rojas,
HAJB 10190.
84 ... Camino & al.
DISTRIBUTION: In the insular Caribbean, this species has been recorded from Cuba,
Dominica, Jamaica, Puerto Rico and Trinidad & Tobago (Minter et al. 2002).
COMMENTS -— This species is frequent in Cuba. After studying type material
of Diderma cubense, we agree with Farr (1976) that this name is a synonym of
D. effusum and not of D. testaceum, as Martin & Alexopoulos (1969) had
indicated previously (Camino et al. 2005).
* Diderma testaceum (Schrad.) Pers. Syn. Meth. Fung.: 167 (1801).
MATERIAL EXAMINED. LOCALITY 11.15, 25-XI-2002, leg. M. Camino, HAJB 9430.
DisTRIBUTION: In the insular Caribbean, this species has been recorded from Cuba,
Guadeloupe, Jamaica and the Dominican Republic (Minter et al. 2002).
ComMENTSs - This species is characterized by sessile sporocarps which are
grouped, pulvinate, and whitish in colour. The peridium is double with the
outer layer porcelain-like, and the columella is prominent and hemispherical.
The spores are 8-10 um diam., pale brown by transmitted light, and slightly
ornamented. The colour of the sporocarps of our samples was almost white, not
pinkish as usually described for this species, but other characters were similar.
* Didymium bahiense Gottsb., Nova Hedwigia 15: 365 (1968). Fics. 4-6
MATERIAL EXAMINED. LOCALITY 8.2.7, on branches of Thunbergia grandiflora, 30-V-
90, leg. M. Benitez & M. Camino, HAJB 6657. Locatity 13.1, on leaves and branches,
20-XI-2003, leg. M. Camino, J.L. Ortiz & J.M. Pérez, HAJB 9827. Loca.iry 2.1, on dead
leaves, 15-XII-2003, leg. M. Camino, M. Cabarroi & J.M. Pérez, HAJB 9900.
DISTRIBUTION: These samples are the first records from Cuba and the insular
Caribbean.
ComMENTSs -— The specimens studied fruited on remains of twigs and leaves, and
show characteristics typical for this species. These include a white, lenticular
to subglobose columella, a transparent peridium covered with crystals of
calcium carbonate and spores 9-10 um diam., with groups of larger and darker
warts. This species is has been recorded previously from Brazil, Ecuador and
Venezuela (Farr 1976).
Didymium clavus (Alb. & Schwein.) Rabenh., Deutschl. Krypt.-Fl. 1: 280 (1844).
MATERIAL EXAMINED. LOCALITY 8.2.2, on twigs of Thunbergia sp., 22-V-96, leg. M.
Camino & O. Alvarez, HAJB 7532. Loca.ity 20.1, on wet rotten wood, 07-VII-2004,
leg. M. Camino, HAJB 10408.
DISTRIBUTION: Didymium clavus is abundant in the insular Caribbean (Minter et al.
2001).
COMMENTS - This species is easy to distinguish because of the black base of
the discoid sporotheca and the dark stipe. The specimens studied had capillitia
with dark forked or three-forked tips and spores 6-7 um diam., with small
warts and some groups of larger and darker warts.
Myxomycete biota of Cuba (1) ... 85
* Didymium difforme (Pers.) Gray, Nat. Arr. Brit. Pl. 1: 571 (1821).
MATERIAL EXAMINED. LOCALITY 11.9, on leaves, 20-II-00, leg. E. Bécquer, J. Calvifio,
M. Camino, J.M. Pérez, D. Minter & Y. Tykhonenko, HAJB 8653.
DisTRIBUTION: This is the first record from Cuba. In the insular Caribbean, this species
has been recorded previously from Dominica, Haiti and Jamaica (Minter et al. 2002).
COMMENTS — The Cuban collection has characteristics typical of this species:
sessile and depressed sporocarps and rarely plasmodiocarps, without columellae,
a smooth peridium and 8-10 um diam. dark spores with minute warts.
* Didymium nigripes (Link) Fr., Syst. Mycol. 3:119 (1829).
MATERIAL EXAMINED. LOCALITY 14.1, on litter and rotten bark, 13-V-1988, leg. M.
Camino, HAJB 5781. Locatity 11.18, on dry leaf, 26-I-97, HAJB 7637. Locairy 11.9,
on leaves, 20-H-00, leg. E. Bécquer, J. Calvifio, M. Camino, J.M. Pérez, D. Minter & Y.
Tykhonenko, HAJB 8649. Locatiry 8.2.7, on soil and twigs of Delonix regia, 25--2003,
leg. M. Clavel, HAJB 9520. Loca.iry 13.6, on dead leaf of Psychotria, 18-XI-2003, leg.
M. Camino, J.L. Ortiz & J.M. Pérez, HAJB 9764. LocALity 11.19, on bark of Samanea
saman, 1-2004, leg. M. Camino, J.L. Ortiz, HAJB 9629.
DisTRIBUTION: ‘These are the first records from Cuba. In the insular Caribbean, this
species has been recorded previously from the American Virgin Islands, Antigua,
Dominica, the Dominican Republic, Haiti, Jamaica, Puerto Rico and Trinidad & Tobago
(Minter et al. 2002).
ComMENTs - This species is frequent on twigs and leaf remains. Didymium
nigripes is close to D. bahiense but is easily differentiated by its blackish stipe,
areolate peridium and smaller, biconvex and dark columella.
Didymium squamulosum (Alb. & Schwein.) Fr., Symb. Gasteromyc.: 19 (1817).
= Didymium costatum Fr., Syst. Mycol. 3: 118 (1829).
MATERIAL EXAMINED. LOCALITY 8.2.4, on stump of tree, 31-I-1991, leg. M. Benitez &
D. Lopez, HAJB 6688.
DISTRIBUTION: This cosmopolitan species was noted (as Didymium costatum) from Cuba
by Berkeley (1869), Massee (1892) and Camino et al. (2007).In the insular Caribbean, it
has been recorded from Cuba, Dominica, the island of Hispaniola, Jamaica, Martinique,
Puerto Rico and Trinidad & Tobago (Minter et al. 2002).
* Echinostelium minutum de Bary, in Rostafinski, Sluzowce Monogr.: 215 (1874).
MATERIAL EXAMINED. LOCALITY 4.4, in moist chamber culture on bark of Pinus sp.,
DWM 7050bc. Locauity 4.4., on Quercus sp. in moist chamber culture, DWM 705 bc.
Locauity 11.3, on bark of Pinus caribaea moist chamber culture, 25-XI-02, leg. S.
Stephenson (no specimen) DWM 6769bc. Locauity 8.2.7, on bark of Diospyros discolor
in moist chamber culture, 12-IH-2003, leg. M. Camino DWM 6807be.
DISTRIBUTION: This is a cosmopolitan corticolous species. Together with specimens
from the Ecological Reserve “Alturas de Banao” (Camino et al. 2007), these are the
first records for Cuba. In the insular Caribbean, this species has been recorded from
Dominica, Guadeloupe, Grenada, Jamaica, and Trinidad & Tobago (Minter et al. 2002).
86 ... Camino & al.
Hemitrichia calyculata (Speg.) M.L. Farr, Mycologia 66(5): 887 (1974). Fias. 7-9
MATERIAL EXAMINED. LOCALITY 1.1, on dead humid trunk, 20-VI-2000, leg. M.
Camino, HAJB 8733. Loca.ity 1.2, on dead wood, 16—XH-2003, leg. M. Cabarroi, M.
Camino & J.M. Pérez, HAJB 9946. Loca.ity 11.20, on rotten wood, 27-I-2004, leg. M.
Camino, M. Cabarroi & J.M. Pérez, HAJB 9963, HAJB 9976. LocALity 10.1, on rotten,
wooden remains, 8-IV-2004, leg. G. Arnold, M. Camino, S$. Maldonado, J.M. Pérez &
G. Recio, HAJB 10275. Loca.ity 19.1, on dead trunk, 03-VII-2004, leg. M. Camino,
HAJB 10342.
DISTRIBUTION: This is one of the most frequent species in Cuba and is mentioned in
several earlier works (Berkeley 1869, Massee 1892) and more recently by Farr (1976),
Camino (1993, 1998b), Pérez & Camino (2000), Camino & Pérez (2001), and Camino
et al. (2005, 2007). In the insular Caribbean, it has been recorded from Cuba, Dominica,
the Dominican Republic, Guadeloupe, Jamaica, Puerto Rico and Trinidad & Tobago
(Minter et al. 2002).
CoMMENTS — The sample HAJB 10275 has a cylindrical, blackish and very
slighty developed stipe about 1 mm long and a capillitium with 6-8 um diam.
elaters with few obtuse free ends. The spores are 7-8 um diam., subreticulate.
Hemitrichia serpula (Scop.) Rostaf. & Lister, Monogr. Mycetozoa: 179 (1894).
[utecd (UEIY,
MATERIAL EXAMINED. LOCALITY 11.9, on rotten petiole of Roystonea regia, 20-[-00,
leg. E. Bécquer, J. Calvifio, M. Camino, J.M. Pérez, D. Minter & Y. Tykhonenko, HAJB
8657. LOCALITY 11.20, on rotten wood, 27-I-2004, leg. M. Camino, M. Cabarroi & J.M.
Pérez, HAJB 9976. Ibidem, on rotten petiole of Roystonea regia, HAJB 9977. LOCALITY
11.5, on rotten petiole of Roystonea regia, 28-I-2004, leg. M. Cabarroi, M. Camino,
J.M. Pérez &. T. Rojas, HAJB 10040. Locauity 11.1, on soil, 29-I-2004, HAJB 10108.
Loca.iry 11.14, on rotten wood, 28-I-2004, leg. M. Camino, HAJB 10187.
DIsTRIBUTION: This is a very frequent species in Cuba. It has been mentioned by Berkeley
(1869), Massee (1892), Farr (1976), Camino (1993, 1998b), Pérez & Camino (2000),
Camino & Pérez (2001), and Camino et al. (2005, 2007). In the insular Caribbean, it has
been recorded from Cuba, Dominica, the Dominican Republic, Guadeloupe, Jamaica,
Puerto Rico, Saint Lucia and Trinidad & Tobago (Minter et al. 2002).
Lamproderma arcyrionema Rostaf,, Sluzowce Monogr.: 208 (1874).
MATERIAL EXAMINED. LOCALITY 11.7, on dead trunk, 20-III-1999, leg. M. Camino, D.
Minter & J.M. Pérez, HAJB 8439. Loca.ity 11.21, on decayed wood, 19-VII-2003,
leg. M. Camino, HAJB 9600. Loca.ity 11.9, on rotten wood of Samanea saman, 14-
X1-2003, leg. M. Camino & J.L Ortiz, HAJB 9639. Locauity 13.3, on wooden remains,
21-XI-2003, leg. M. Camino, J.L. Ortiz & J.M. Pérez, HAJB 9846. Loca.ity 20.1, on
dead trunk, 07-VIH-2004, leg. M. Camino, HAJB 10406.
Fics. 7-9 Hemitrichia calyculata HAJB 9946. 7. Capillitium (bar = 5 um). 8. Spore (bar = 2 um).
9. Detail of spore ornamentation (bar = 1 um). Fics. 10-12 Hemitrichia serpula HAJB 9977.
10. Capillitium (bar = 5 um). 11. Spore (bar = 2 um). 12. Detail of spore ornamentation HAJB
10187 (bar = 1 um). Fics. 13-14. Stemonitis axifera HAJB 9916. 13. Spore (bar = 1 ttm). 14. Detail
of spore ornamentation (bar = 1 um).
Myxomycete biota of Cuba (1) ... 87
88 ... Camino & al.
DISTRIBUTION: It is frequently found in Cuba on ligneous remains. The species was cited
recently by Camino & Pérez (2001) and Camino et al. (2007). In the insular Caribbean,
it has been recorded from Cuba, Dominica, the island of Hispaniola (the Dominican
Republic and Haiti), Jamaica, Puerto Rico and Saint Lucia (Minter et al. 2002).
CoMMENTSs -— This species is well characterized by globose sporothecae, a silvery
to bronze peridium, a columella tapering towards the centre of the sporotheca
and a capillitium with main branches and sinuous filaments. The spores are
(6-) 7-9 um diam., with groups of darker warts.
Licea biforis Morgan, J. Cincinnati Soc. Nat. Hist. 15: 131 (1893).
MATERIAL EXAMINED. LOCALITY 8.2.8, on litter of Thunbergia grandiflora, 14-X-1992,
leg. W. Alonso & M. Camino, HAJB 7050. Ibidem, on bark of Pouteria mammosa in
moist chamber culture, 12-III-2003, leg. M. Camino, obtained by D.W. Mitchell, DWM
6830bc. Loca.ity 4.2, on bark, 25-II-1997, leg. C. Panfet, HAJB 7673.
DISTRIBUTION: Samples HAJB 7050 and 7673 were mentioned by Minter et al. (2001).
In the insular Caribbean, this species has been recorded from Cuba and Jamaica (Minter
et al. 2002).
ComMENTs - This species is characterized by its elongated to sinuous, laterally
flattened plasmodiocarps. The colour is very variable, ranging from white
to fuscous depending on the material incorporated from the substratum.
Dehiscence is by a pale longitudinal slit. It is found on bark and litter and,
because of its small size, usually observed when studying other samples by
OM.
* Licea pedicellata (H.C. Gilbert) H.C. Gilbert, in Martin, Mycologia 34(6): 702
(1942).
MATERIAL EXAMINED. LOCALITY 8.2.7, in moist chamber culture on bark of Terminalia
eriostachya, 12-III-2003, leg. M. Camino, obtained by D.W. Mitchell, DWM 6842bc.
DISTRIBUTION: This is a new record from Cuba. In the insular Caribbean, this species
has been recorded from Grenada and Puerto Rico (Minter et al. 2002).
CoMMENTS - This corticolous species is distinguished by its stout furrowed
stalk and is the only common stipitate Licea that dehisces by peridial platelets.
The spores have a paler region and appear smooth or minutely spinulose by
transmitted light.
* Licea pseudoconica T.E. Brooks & H.W. Keller, in Keller & Brooks, Mycologia
69(4): 678 (1977).
MATERIAL EXAMINED. LOCALITY 8.2.7, in moist chamber culture on bark of Antidesma
montanum, 12-III-2003, leg. M. Camino, obtained by D.W. Mitchell, DWM 6821 bc.
DISTRIBUTION: This is a new record from Cuba and the insular Caribbean.
COMMENTS —- A corticolous species that, at first sight, resembles a small
pyrenomycete because its sessile sporocarps have a conical apical deposit of
Myxomycete biota of Cuba (1) ... 89
refuse material. Dehiscence is by rupture of the lateral wall, after which the basal
part persists as a shallow cup. The spore-mass is dark brown to almost black.
The spores are 9.5-11 (-13) um diam., olivaceous brown, globose, smooth and
with a thin, pale germination pore.
Lycogala epidendrum (L.) Fr., Syst. Mycol. 3: 80 (1829).
MATERIAL EXAMINED. LOCALITY 8.2.3, on wood of Pinus sp., 11-VI-2001, leg. M.
Clavel, HAJB 8819. Loca.iry 11.20, on decayed wood, 27-I-2004, leg. M. Camino, M.
Cabarroi & J.M. Pérez, HAJB 9975; Locatity 11.4, on decayed wood, 28-I-2004, leg.
M. Camino, M. Cabarroi, J.M. Pérez & T. Rojas, HAJB 10036; Locaity 11.1, on rotten
petiole of Roystonea regia, 28-1-2004, leg. M. Camino, M. Cabarroi, A. Fidalgo, J.M.
Pérez & T. Rojas, HAJB 10106.
DIsTRIBUTION: ‘This species is very frequent in Cuba and other Caribbean countries. It
was one of the first six myxomycetes recorded from the island (Montagne 1845). It has
been cited in early (Berkeley 1869, Massee 1892) and more recent works (Farr 1976,
Camino 1998b, Camino & Pérez 2001, Camino et al. 2005, Perez & Camino 2000). In the
insular Caribbean, it has been recorded from Cuba, Dominica, the Dominican Republic,
Guadeloupe, Grenada, Jamaica, Puerto Rico, Saint Lucia and Trinidad & Tobago (Minter
et al. 2002).
* Macbrideola cornea (G. Lister & Cran) Alexop., Mycologia 59(1): 112 (1967).
MATERIAL EXAMINED. LOCALITY 8.2.7, on bark of Antidesma montanum bark in moist
chamber culture, 12-III-2003, leg. M. Camino, DWM 6820bc (slide only),
DISTRIBUTION: This is a new record from Cuba and the insular Caribbean.
CoMMENTS - This is a corticolous species usually found on bryophytes. The
distinctly hollow stalk is red to yellow at the base and tapers upwards and extends
into about the centre of the sporotheca to form the columella. The spherical
sporothecae are mostly 0.1-0.3 mm diam. The peridium is partly persistent
as a collar around the stalk apex. The capillitium is dark brown, branched
dichotomously, usually only 1-3 times, tapered outwards but remaining fairly
stout at the periphery. The spore—mass is dark brown while the spores are lilac-
brown or grey-brown, verrucose, and 8-9 «tm diam.
Perichaena chrysosperma (Curr.) Lister, Monogr. Mycetozoa: 196 (1894).
MATERIAL EXAMINED. LOCALITY 11.1, on decayed wood, 28-I-2004, leg. M. Camino,
M. Cabarroi, A. Fidalgo, J.M. Pérez & T. Rojas, HAJB 10110. Loca.iry 10.1, on rotten
wood, 08-IV-2004, leg. G. Arnold, M. Camino, S. Maldonado, J.M. Pérez & G. Recio,
HAJB 10276.
DISTRIBUTION: The species has been cited for Cuba in several works (Berkeley 1869,
Massee 1892, Farr 1976, Camino, 1993, 1998b, Camino & Perez 2001, Camino et al.
2005, 2007), and is widely distributed in the insular Caribbean, having been recorded
from Cuba, Dominica, the Dominican Republic, Granada, Guadeloupe, Jamaica, Puerto
Rico, Saint Lucia and Trinidad & Tobago (Minter et al. 2002).
90 ... Camino & al.
ComMENTs ~ This material exhibits characteristics typical of the species: short,
elongated or reniform plasmodiocarps, threads of the capillitium with scarce
and prominent spines up to 4 um long and spores 9-11 um diam.
* Physarum decipiens M. A. Curtis, Amer. J. Sci. Arts 6: 352 (1848).
MATERIAL EXAMINED. LOCALITY 8.2.2, 100 msm, on twigs of Thunbergia sp., 22-V-
1996, leg. M. Camino & O. Alvarez, HAJB 7531 and HAJB 7532.
DISTRIBUTION: This species was not cited for the insular Caribbean by Minter et al. (2001,
2002) and Farr (1976) mentioned it for the Neotropics from Bolivia and, doubtfully,
Costa Rica. This is therefore a new record from Cuba and the insular Caribbean.
COMMENTS — Physarum decipiens is a species very close to P. auriscalpium
Cooke and P. serpula Morgan in form. According to Farr (1961), they differ
in habit of development and structure of the peridial calcium carbonate.
Physarum decipiens has sessile sporocarps, or short plasmodiocarps, with a
typical “badhamioid” capillitium formed by pronounced, yellowish nodes of
calcium carbonate. A good photograph of this species can be seen in Neubert
et al. (1995).
* Physarum flavicomum Berk., London J. Bot. 4: 66 (1845).
MATERIAL EXAMINED. LOCALITY 11.16, on bark of rotten trunk, 20-IX-1997, leg. U.
Eliasson, M. Camino, E. Bécquer, & J. Calvifio, HAJB 7894.
DISTRIBUTION: This species appears to be uncommon in the insular Caribbean and
was cited only from Antigua and Trinidad & Tobago by Minter et al. (2001, 2002). This
collection is the first record of this species from Cuba.
CoMMENTS - This species is characterized by the following features: stipitate
sporocarps; usually globose, nodding sporothecae, 0.3-0.4 mm diam.; very
delicate, generally iridescent peridia, with whitish to yellowish deposits of
calcium carbonate; stalks up to 2 mm long, thin, yellowish to paler towards the
apex, dark reddish towards the base; capillitia with white to yellowish nodes.
The spores are brown to pale purple, 8-10 um diam, densely warted, with
groups of more prominent warts.
* Physarum globuliferum (Bull.) Pers., Syn. Meth. Fung.: 175 (1801).
MATERIAL EXAMINED. LOCALITY 11.22, on wooden remains of Samanea saman, 20-XI-
2001, leg. E. Bécquer, M. Camino, D. Minter & J.M. Pérez, HAJB 8948a, Locauiry 4.1,
on rotten palm petiole, 09—XI-2002, leg. M. Camino & J.L. Ortiz, HAJB 9222.
DISTRIBUTION: This collection represents the first record from Cuba but, in the insular
Caribbean, this species has been recorded previously from Guadeloupe, Jamaica and
Trinidad & Tobago (Minter et al. 2001, 2002).
COMMENTS ~ This species is characterized by its whitish stalked fructifications,
globose sporothecae and whitish peridium. The stalk is calcareous, fragile,
narrower towards the apex, and more than four times as long as the diameter of
Myxomycete biota of Cuba (1) ... 91
the sporotheca. The capillitium is hyaline with small whitish nodes. The spores
are subglobose, 7-10 um diam., verruculose with more prominent groups of
warts.
* Physarum gyrosum Rostaf., Sluzowce Monogr.: 111 (1874).
MATERIAL EXAMINED. LOCALITY 8.2.1, on leaves, I-2004, leg. M. Cabarroi, HAJB 9952.
DISTRIBUTION: This is a new record from Cuba and the insular Caribbean.
ComMENTs - This species is characterized by its crowded and sinuous aethalial
fructifications. The straw-coloured stipe is a continuation of the hypothallus.
The peridium is thin, membranous, whitish to cream to straw-coloured. The
elastic capillitium is dense with abundant very elongated and whitish fusiform
nodes. The spores are 7-10 um diam., with spinulose ornamentation.
* Physarum mutabile (Rostaf.) G.Lister, in Lister, Monogr. Mycetozoa, ed. 2: 53
C1915):
MATERIAL EXAMINED. LOCALITY 11.4, on rotten bark of Jatropha curcas, 28-1-2004, leg.
M. Camino, M. Cabarroi, A. Fidalgo, J.M. Pérez & T. Rojas, HAJB 10038.
DIsTRIBUTION: Physarum mutabile appears to be rare in the Neotropics (Farr 1976), and
this is a new record from Cuba and the insular Caribbean.
COMMENTS - The Cuban material has subglobose, crowded, white to beige
sporocarps with a thick peridium and a reticulated capillitium with scanty,
small and whitish nodes. At 10-12 tm diam., the spores are larger than those
(8-10 um), cited by Martin & Alexopoulos (1969).
* Physarum nicaraguense T. Macbr., Bull. lowa Univ. Lab. Nat. Hist. 2: 382 (1893).
MATERIAL EXAMINED. LOCALITY 8.1.1, on trunk of Erythrina grisebachii, 12-XU-1994,
leg. M. Clavel, HAJB 7312.
DIsTRIBUTION: This collection represents the first record for Cuba, but this species has
been recorded previously in the insular Caribbean from Haiti, Jamaica, Puerto Rico and
Trinidad & Tobago (Minter et al. 2001, 2002).
CoMMENTS — The Cuban sample has abundant, crowded fructifications, with
subglobose to sinuous whitish sporothecae covered by calcium carbonate, a
short, blackish stipe and a blackish hypothallus. The capillitium is dense with
whitish broad and angular nodes that join in the centre forming a typical
whitish, squashed pseudocolumella. The spores are 9-11 tm diam., dark
purplish and warted.
Physarum nucleatum Rex, Proc. Acad. Nat. Sci. Philadelphia 43: 389 (1891).
MATERIAL EXAMINED. LOCALITY. 11.13, on bark, 19-IX-1997, leg. U. Eliasson, M.
Camino, E. Bécquer & J. Calvifio, HAJB 7843 and HAJB 7844.
92 ... Camino &al.
DISTRIBUTION: This species was cited from Cuba for the first time by Minter et al. (2001)
and there are two new collections. In the insular Caribbean it has also been recorded
from Dominica, Jamaica, Puerto Rico, and Trinidad & Tobago (Minter et al. 2002).
COMMENTS ~ This species has stipitate sporocarps with whitish globose, fragile
sporothecae, 0.5-0.9 mm diam. The stipe is long, 2-3 times the diameter of the
sporotheca, straw-yellow, without dark granulations, flattened and expanded
towards the base. The reticulate capillitium has small whitish nodes that
sometimes join in the centre as a globose whitish pseudocolumella (HAJB
7843). The spores are 6-7.5 um diam., pale purplish by OM, and warted with
slightly darker areas. —
* Physarum oblatum T. Macbr., Bull. Iowa Univ. Lab. Nat. Hist. 2: 384 (1893).
MATERIAL EXAMINED. LOCALITY 2.1, on rattan of Cissus sicyoides, 15—XII-2003, leg. M.
Cabarroi, M. Camino & J.M. Pérez, HAJB 9898.
DIsTRIBUTION: This is a new record from Cuba. In the insular Caribbean, this species
has been recorded from Dominica and Jamaica (Minter et al. 2001, 2002).
CoMMENTS - This species is distinguished by its stipitate sporocarps with 0.4-0.7
mm diam. lenticular to subglobular sporothecae. It has a thick, but very fragile,
peridium, covered with crystals of calcium carbonate that give it a very warted
appearance. It is orange yellow in colour. The stalk is of similar length to the
diameter of the sporotheca, rigid, and from reddish brown to blackish red.
The capillitium is reticulate with abundant yellowish to orange polyhedric,
calcareous to fusiform nodes. The spores are globose, 12-13 um diam., minutely
warted and with groups of darker warts.
Physarum pusillum (Berk. & M. A. Curtis) G. Lister, in Lister, Monogr. Mycetozoa,
ed. 2: 64 (1919).
MATERIAL EXAMINED. LOCALITY 15.1, on dry leaf, 01-XII-1994, leg. M. Bénitez, HAJB
DIAN
DIsTRIBUTION: It has been cited previously from Cuba by Camino & Pérez (2001) and
Camino et al. (2007). In the insular Caribbean, it has also been recorded from Dominica,
the island of Hispaniola (the Dominican Republic and Haiti), Jamaica, and Puerto Rico
(Minter et al. 2002).
ComMENTs - Physarum pusillum is characterized by its whitish and lenticular
sporothecae with irregular dehiscence and blackish stipes.
* Physarum rigidum (G. Lister) G. Lister, in Lister, Monogr. Mycetozoa, ed. 3: 36 (1925).
MATERIAL EXAMINED. LOCALITY 6.1, on woody remains, 19-VIII-1987, leg. S.
Maldonado, R. Rankin & H.D. Gomez, HAJB 5434. Locatity 9.1, on decayed wood,
VIII-1993, leg. H. Notario, HAJB 7108.
DISTRIBUTION: This is a new record from Cuba. In the insular Caribbean, this species
has been recorded from the American Virgin Island, Jamaica, Puerto Rico and Trinidad
& Tobago (Minter et al. 2001, 2002).
Myxomycete biota of Cuba (1) ... 93
COMMENTS ~— This species is characterized by its stipitate sporocarps with very
fragile white lenticular sporothecae, approximately 0.7-1 x 0.3-0.4 mm. The
stipe is 1.8-2.2 mm long, straw yellow, more orange to orange brown in the
lower part, tapered towards the apex, and without lime or granular material.
The capillitium has a “badhamioid” structure, formed by parallel yellowish
threads from the apex to the base of the sporotheca, with yellowish fusiform
elongated nodes. The spores are 8-10 um diam. and warted.
* Physarum roseum Berk. & Broome, J. Linn. Soc., Bot. 14 (2): 84 (1873).
MATERIAL EXAMINED. LOCALITY 3.1, on trunk of Syzygium jambos, 5-X1I-2001, leg. M.
Camino & J.M. Pérez HAJB 9013.
DIsTRIBUTION: This is a new record for Cuba. In the insular Caribbean this species was
previously recorded from Dominica and Jamaica (Minter et al. 2002).
COMMENTS — Physarum roseum is characterized by its stipitate and grouped,
globose sporocarps, fragile sporothecae, pink to red purple or red scarlet,
0.5-0.6 mm diam., and with irregular to petaloid dehiscence. The stalk is dark
brown, 1-1.2 mm long, without calcium carbonate. The capillitium is formed
by a net of filaments with large angular reddish to purple nodes, according to
drawings by Lister (1925) and Emoto (1977).
* Physarum stellatum (Massee) G.W. Martin, Mycologia 39(4): 461 (1947).
MATERIAL EXAMINED. LOCALITY 4.3, on rotten wood, 12—XI-2004, leg. M. Camino, M.
Cabarroi, L. Castillo, J.M. Pérez, K. Salas, HAJB 10552. LOCALITY 11.10, on bark of rotten
trunk, 27-I-1997, leg. M. Camino, HAJB 7643; Loca.ity 11.2, on rotten stalk of Piper
auritum, 21-IX-1997, leg. U. Eliasson, M. Camino, E. Bécquer & J. Calvino, HAJB 7913;
Loca.iry 11.17, on rotten rachis of Roystonea regia, 14—X1-2003, leg. M. Camino & J.L.
Ortiz, HAJB 9635. Locatirty 1.2, in trunk, 13-I-2002, leg. J.M. Pérez & M. Rodriguez.,
HAJB 9121. Loca.ity 11.22, on wooden remains of Samanea saman, 20-XI-2001, leg.
E. Bécquer, M. Camino, D.W. Minter & J.M. Pérez, HAJB 8948b.
DISTRIBUTION: This is a new record from Cuba. In the insular Caribbean this species
has been recorded from Dominica, Guadeloupe, Jamaica, Puerto Rico and Trinidad &
Tobago (Minter et al. 2002).
COMMENTS - This species is characterized by the following features: a
calcareous stipe, and a peridium with thick patches or scales of calcium
carbonate and floriform dehiscence into typical elongated lobes; a globose,
white pseudocolumella in the centre of the sporotheca; a capillitium with few,
whitish and rounded nodes; globose spores, 8.5-9.5 um diam., pale purplish,
minutely spinose or warted, with warts distributed homogeneously.
Excellent colour illustrations of this species can be found in the books by
Emoto (1977) and Lister (1925) and, more recently, in the photographs of
Neubert et al. (1995).
94 ... Camino &al.
Physarum viride (Bull.) Pers., Ann. Bot. (Usteri) 15: 6 (1795).
MATERIAL EXAMINED. LOCALITY 16.1, in trunk, 01-05-1989, leg. M. Benitez & H.D.
Gomez, HAJB 6291. Locauity 13.3, on trunk, 21-XI-2003, leg. M. Camino, J.L. Ortiz
& J.M. Pérez, HAJB 9836.
DISTRIBUTION: This species has been recorded from Cuba by Camino & Pérez (2001)
(Sancti Spiritus) and Camino et al. (2007). In the insular Caribbean, it has also been
recorded from Dominica, Guadeloupe, Jamaica, Puerto Rico and Trinidad & Tobago
(Minter et al. 2002).
COMMENTS -— Physarum viride is characterized by its lenticular and usually
nodding sporothecae, a greenish to orange peridium that breaks into petaloid
lobes, and a dark stalk, paler at the apex. It has a hyaline, reticulate capillitium
with yellowish calcareous nodes. The spores are 7-8 um diam., globose to
subglobose, warted, with groups of darker warts.
* Reticularia jurana Meyl., Bull. Soc. Vaud. Sci. Nat. 44: 297 (1908).
MATERIAL EXAMINED. LOcALIry 4.3, on rotten wood, 12-XI-2004, leg. M. Camino, M.
Cabarroi, L. Castillo, J.M. Pérez, K. Salas, HAJB 10552.
DISTRIBUTION: This collection represents the first record from Cuba and the insular
Caribbean.
ComMENTS - This species forms a pulvinate brownish aethalium, and a
capillitium with filaments that form an isodiametric reticulum, where its
threads turn out to be perforated. The spores are 6-8 um diam., and reticulated
over more or less than half of the surface of spore.
Stemonitis axifera (Bull.) T. Macbr., N. Amer. Slime-Moulds: 120 (1899). Fias. 13-14
MATERIAL EXAMINED. LocALiry 11.17, on bark of Samanea saman, 29-I-1997, leg. M.
Camino, HAJB 9629. Locatity 11.6, on rotten wood of Guarea guara, 24-—XI-2002, leg.
M. Camino, HAJB 9410. Locatirty 1.2, on rotten trunk, 15-XII-2003, leg. M. Cabarroi,
M. Camino & J.M. Pérez, HAJB 9916. LocALiry 8.2.5, on trunk of Pinus sp., 22-X-2004,
leg. M. Camino, HAJB 10475.
DIsTRIBUTION: This species is frequent in Cuba and was cited by Camino (1998a) and
Camino et al. (2003). In the insular Caribbean it also has been recorded from Dominica,
Guadeloupe, the island of Hispaniola (the Dominican Republic and Haiti), Jamaica,
Martinique and Trinidad & Tobago (Minter et al. 2002).
COMMENTS ~ This species is frequent in Cuba. It is characterized by its small
spores 5~7 um diam., smooth by transmitted light, but with a lax and irregular
ornamentation formed by baculae and a very thin net observable by SEM.
Stemonitis fusca Roth, Bot. Mag. (R6mer & Usteri) 1(2): 26 (1787).
MATERIAL EXAMINED. LOCALITY 11.19, on bark of Tabebuia sp., 14-XI-2003, leg. M.
Camino & J.L. Ortiz, HAJB 9637; Locauity 11.20, on trunk of Tradescantia sp., 27-I-
2004, leg. M. Camino, M. Cabarroi & J.M. Pérez, HAJB 9965a and HAJB 9965b; LOCALITY
11.1, on rotten trunk of Cecropia schreberiana, 28-I-2004, leg. M. Camino, M. Cabarroi,
Myxomycete biota of Cuba (1)... 95
A. Fidalgo, J.M. Pérez & T. Rojas, HAJB 10109. Loca.ity 13.3, on trunk, 21-XI-2003,
leg. M. Camino, J.L. Ortiz & J.M. Pérez, HAJB 9839. LocALity 1.2, rotten trunk, 16—XII-
2003, leg. M. Cabarroi, M. Camino & J.M. Pérez HAJB 9922. Loca ity 10.1, on petiole
of Roystonea regia, 08-IV-2004, leg. G. Arnold, M. Camino, S. Maldonado, J.M. Pérez
& G. Recio, HAJB 10278.
DIsTRIBUTION: This species is one of the most frequent of the genus Stemonitis in Cuba.
It was cited by Berkeley (1869), Massee (1892), Farr (1976), Camino (1998a), Camino
& Pérez (2001), and Camino et al. (2003, 2005). In the insular Caribbean it has been
recorded from Cuba, Dominica, the Dominican Republic, Guadeloupe, Jamaica, Puerto
Rico and Trinidad & Tobago (Minter et al. 2002).
Stemonitis herbatica Peck, Annual Rep. New York State Mus. 26: 75 (1874).
MATERIAL EXAMINED. Locatity 18.1, on trunk of Pinus cubensis, 1-V1I-2004, leg. M.
Camino, HAJB 10324; Ibidem, on bark of trunk of Pinus cubensis, HAJB 10325.
DISTRIBUTION: This species was cited from Cuba by Camino (1998a) and Camino
et al. (2003). In the insular Caribbean it has also been recorded from Dominica, the
Dominican Republic, Guadeloupe, Jamaica, Martinique and Puerto Rico (Minter et al.
2002).
ComMENTS - Both collections have sporocarps up to 1 mm long and a short
stalk (1/5 the total height). The capillitium is formed by perpendicular branches
as shown by Nannenga-Bremekamp (1974) and Ing (1999), with small and thin
meshes, 3-25 um diam., and small spines. The spores are globose, 7-9 um diam.,
purplish and with regularly distributed minute warts. Although this species
typically occurs on herbaceous substrata (leaves, stems and debris), lignicolous
records are also known (Martin & Alexopoulos 1969, Camino 1998a).
Stemonitis splendens Rostaf., Sluzowce Monogr.: 195 (1874).
MATERIAL EXAMINED. LOCALITY 11.21, on leaf of Guarea guidonia, 15-X1-2003, leg. M.
Camino & J.L. Ortiz, HAJB 9711; Loca.ity 11.12, on bark of Cordia sp., 8-VIII-2004,
leg. M. Camino, HAJB 10694. Loca tity 13.3, on trunk, 21-XI-2003, leg. M. Camino,
J.L. Ortiz & J.M. Pérez, HAJB 9838. LOCALITY 8.2.6, on trunk of Hibiscus elatus and
leaves of Poaceae, 21-XI-2003, leg. M. Clavel, HAJB 9951.
DIsTRIBUTION: This species is abundant and was cited from Cuba by Lister (1894,
1925), Macbride & Martin (1934), Martin (1949), Farr (1976), Camino (1998a, 1998b),
Pérez & Camino (2000), Camino & Pérez (2001), and Camino et al. (2003, 2005). In the
insular Caribbean it has been recorded from Cuba, Dominica, Guadeloupe, Jamaica,
Martinique, Puerto Rico and Trinidad & Tobago (Minter et al. 2002).
Stemonitopsis typhina (FH. Wigg.) Nann.-Bremek., Nederlandse
Myxomyceten: 209 (1975) [“1974”]. Figs. 15-16
= Stemonitis typhoides (Bull.) DC., Fl. Frang. 2: 257 (1805).
MATERIAL EXAMINED. LOCALITY 11.14, on rotten trunk of Cinnamomum sp., 30-
I-2004, leg. M. Camino, M. Cabarroi, A. Fidalgo, J.M. Pérez & T. Rojas, HAJB 10192.
Locauity 19.1, on trunk, 3-VII-2004, leg. M. Camino, HAJB 10349, HAJB 10352 and
HAJB 10353.
96 ... Camino & al.
DISTRIBUTION: Stemonitopsis typhina has been cited from Cuba by Montagne (1845),
Berkeley (1869), Massee (1892), Farr (1976), Camino (1998a), Camino & Pérez (2001)
and Camino et al. (2003). In the insular Caribbean, this species has been recorded from
Cuba, Dominica, the Dominican Republic, Guadeloupe, Jamaica, Puerto Rico and
Trinidad & Tobago (Minter et al. 2002).
ComMENTs — This species is characterized by its sporocarps, which have a silver
peridium and membranous, silvery remains on the stalk. Some Cuban samples,
however, do not posses these features, but the morphology of the sporothecae
and the spores (6-8 tum diam., warted, with scatter larger warts) enabled them
confidently to be identified as this species.
Trichia affinis de Bary, in Fuckel, Nassauischen Vereins Naturk. 23-24: 336 (1870).
MATERIAL EXAMINED. LOCALITY 11.4, on rotten wood, 19-HI-1999, leg. M. Camino, D.
Minter, J.M. Pérez, HAJB 8410. Loca.ity 11.8, on dead trunk of Guarea guara, 21-XI-
2001, leg. E. Bécquer, M. Camino, D. Minter & J.M. Pérez, HAJB 8965. Loca.ity 13.1,
on wet dead wood, 15-III-2006, leg. M. Camino, HAJB 10828.
DIsTRIBUTION: Trichia affinis is cited from Cuba by Camino et al. (2005, 2007). This
species is new for the insular Caribbean, but most records of T. favoginea sensu Farr from
Cuba, Dominica, the Dominican Republic, Jamaica, and Trinidad & Tobago (Minter et
al. 2002) are probably T! affinis.
COMMENTS - This species is characterized by its sessile, crowded, globose to
subglobose sporocarps. The capillitium is formed by yellowish elaters, 5-6 um
diam., without spines, or with very short and dispersed ones. The elaters have
sharp and short free ends. The spores are 10-12 um diam., with approx. 1 um
high reticulations.
Farr (1976) considered Trichia persimilis P. Karst. and T. affinis to be
synonyms of T. favoginea (Batsch) Pers. Farr (1958) mentioned material from
Cuba of T. favoginea deposited in BPI, FH. In our experience, the frequent
species in Cuba is T: affinis whereas T! favoginea and T. persimilis are both very
rare in the island.
* Trichia agaves (G. Moreno, Lizarraga & Illana) Mosquera, Lado, Estrada &
Beltran-Tej., in Lado, Cuad. Trab. Fl. Micol. Iber. 16: 82 (2001). Figs. 17-22
MATERIAL EXAMINED. LOCALITY 13.2, on leaf of Agave perdentata, 21-XI-2003, leg. M.
Camino, J.L. Ortiz & J.M. Pérez, HAJB 9832.
DISTRIBUTION: This collection is a new record for Cuba and the insular Caribbean.
COMMENTS - Trichia agaves is characterized by its sessile, globose to subglobose
sporocarps, 0.5-1 mm diam., grouped to crowded, and often in extensive
groups. The peridium is double with a thick, opaque outer layer which is
Fics. 15-16 Stemonitopsis typhina HAJB 10192. 15. Spore (bar = 2 um). 16. Detail of spore
ornamentation (bar = 1 um). Fics. 17-22 Trichia agaves HAJB 9832. 17-20. Detail of capillitium
(bar = 2 um). 21-22. Spores (bar = 2 pm).
Myxomycete biota of Cuba (1) ... 97
98 ... Camino &al.
ochraceous to deep orange when old. The thin, membranous, translucent, pale
yellow inner layer is attached to the outer layer. Dehiscence is operculate or
sometimes irregular. There is no columella. The capillitium is elastic, yellow by
transmitted light, and orange coloured when mature. The elaters are 2-3 (—4)
uum diam., flexuous, branched, with blunt free ends, and ornamented with few,
1-1.5 um long warts, and very faint spiral bands (visible only by oil immersion).
The spores are yellow in mass, light yellow by transmitted light, (11-)13-15
uum diam., globose to subglobose, warted. With SEM the capillitial elaters are
shown to be ornamented with spirals and the spores with baculae.
This is a species which typically fruits on succulent plants, especially
Crassulaceae and Agave (Liliaceae) and for this reason the epithet “agaves” was
chosen (Moreno et al. 2000).
Tubulifera microsperma (Berk. & M.A.Curtis) Lado, Cuad. Trab. Fl. Micol. Iber. 16:
87 (2001).
= Tubifera microsperma (Berk. & M.A. Curtis) G.W. Martin, Mycologia 39(4): 461 (1947).
= Licea stipitata Berk. & Ravenel, in Berkeley & Curtis, Proc. Amer. Acad. Arts 4: 125 (1860),
nom. illeg., non L. stipitata DC., 1815.
= Tubulina stipitata Berk. & Ravenel ex Rostaf., Sluzowce Monogr.: 223 (1875).
MATERIAL EXAMINED. LOCALITY 17.1, on stump, 4-V-1989, leg. M. Benitez, H.D.
Gomez, H. Saralegui & C. Panfet, HAJB 6346. Locatity 20.1, on rotten bark with
mosses, 7-VII-2004, leg. M. Camino, HAJB 10404.
DISTRIBUTION: This species was cited from Cuba by Berkeley (1869) as Licea stipitata,
by Massee (1892) as Tubulina stipitata, and by Farr (1976), Camino (1998b) and Camino
& Pérez (2001) as Tubifera microsperma. In the insular Caribbean this species has been
reported (as Tubifera microsperma) from Cuba, Dominica, Guadeloupe, Jamaica, Puerto
Rico and Trinidad & Tobago (Minter et al. 2002).
ComMENTSs - This species is easily identified by its pedicelate aethalia. The inside
of the peridium has protuberances in the form of short tubes that resemble the
suckers of octopus tentacles, easily visible by transmitted light and SEM. The
spores are small, with a completely reticulated hemisphere.
Discussion
In this study, 50 species of myxomycetes were identified from 118 specimens
obtained as field collections and from moist chamber cultures in 58 localities
throughout Cuba. Of this total, 25 species are here reported for the first time
from Cuba with 10 representing new records for the insular Caribbean,-
Subreino Neotropical (Takhtajan et al. 1986).
The 50 species represent 12 families of myxomycetes: Arcyriaceae
(Perichaena: 1 species), Clastodermataceae (Clastoderma: 1 _ species),
Cribrariaceae (Cribraria: 4 species), Dianemataceae (Calomyxa: 1 species),
Dictydiaethaliaceae (Dictydiaethalium: 1 species), Didymiaceae (Diderma:
Myxomycete biota of Cuba (1) ... 99
2 species; Didymium: 5 species), Echinosteliaceae (Echinostelium: 1 species),
Liceaceae (Licea: 3 species), Physaraceae (Craterium: 2 species; Physarum: 13
species), Reticulariaceae (Lycogala: 1 species; Reticularia: 1 species; Tubulifera:
1 species), Stemonitidaceae (Comatricha: 2 species; Lamproderma: 1 species;
Macbrideola: 1 species; Stemonitis: 4 species, Stemonitopsis: 1 species) and
Trichiaceae (Hemitrichia: 2 species; Trichia: 2 species). The families that have
the greatest diversity of species are: Physaraceae and Stemonitidaceae, followed
by Didymiaceae, Cribrariaceae, Trichiaceae, Reticulariaceae and Liceaceae. The
remaining families are represented by only one genus and one species.
Species of the Physarales predominate, an aspect already noted by other
authors for tropical and subtropical forests of the American continents
(e.g., Schnittler & Stephenson 2000). Of the 50 studied species most have a
cosmopolitan distribution. Nevertheless Craterium paraguayense, Hemitrichia
calyculata, Physarum flavicomum, P. globuliferum, P. gyrosum, P. nicaraguense,
P. roseum, P. stellatum and Tubulifera microsperma appear to be mainly
Neotropical. In addition, a number of species can be considered as rare or
infrequently noted. These include Comatricha anomala, Cribraria confusa, Licea
pseudoconica, Reticularia jurana and Trichia agaves. In this study, members
of genus Arcyria were not collected, although some, such as A. cinerea and
A. denudata, are very common on fallen wood in tropical forests (Stephenson
et al. 2003). Trichia agaves is a species that develops on succulent plants
(mainly Cactaceae and Agavaceae). Seven species considered herein produced
sporocarps in moist chamber cultures. These were Licea biforis, L. pedicellata,
L. pseudoconica, Calomyxa metallica, Comatricha anomala, Echinostelium
minutum and Macbrideola cornea. In terms of substrata, the foliicolous
species which were recorded in this study were Craterium leucocephalum,
C. paraguayense, Diderma effusum, Didymium bahiense, D. clavus, D. difforme
and D. nigripes. The various species collected from the genus Cribraria occurred
mainly on wood and bark in damp places, often in association with bryophytes.
Collections of Stemonitales and Hemitrichia calyculata were clearly lignicolous.
The frequent occurrence of Hemitrichia serpula on palms is noteworthy, in
particular its occurrence on petioles and rachides of Roystonea regia (Royal
Palm, Arecaceae). Lycogala epidendrum, Physarum globuliferum, P. stellatum
and Stemonitis fusca were also found on these substrata.
Provinces of Cuba with many species are: Ciudad de la Habana, Pinar del
Rio, Sancti Spiritus and Santiago de Cuba. Fewer species have been recorded
from Camagtiey, Guantanamo, Granma, Holguin and La Habana, while there
are no records at all from Matanzas, Villa Clara, Ciego de Avila and Las Tunas.
These are all in or adjacent to the central zone of the island, where the climate
tends to be drier and the vegetation more scanty, factors not very suitable for
growth of myxomycetes. Nevertheless, the central zone also contains the “Sierra
100 ... Camino & al.
de Escambray’, where the climate is less dry, and the vegetation more suitable
for myxomycetes. It would be rewarding to explore that area more fully.
Acknowledgements
This investigation was funded in part by the Research Project of “Ministerio de Asuntos
Exteriores” and “Ministerio de Ciencia y Tecnologia’, Plan Nacional de Investigacion
Cientifica, Desarrollo e Innovacién Tecnolégica REN2002-01965. Another part was
financed through a “Miguel de Cervantes” Scholarship of the University of Alcala, for
the first author, and her field work was supported by the UK Darwin Initiative Projects
Fungi of the Caribbean and Biodiversity Conservation in Cuba. We express our gratitude
to Dasmilia Cruz Arosarena, HAJB reference collection technician, Jardin Botanico
Nacional de Cuba for her help with data relating to material in that collection and J.A.
Pérez and A. Priego “Servicio de Microscopia Electrénica, University of Alcala” for their
invaluable help with the SEM. We wish to thank Dr. Steve Stephenson and Dr. Marcos
Lizarraga for their revision of the manuscript.
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MYCOTAXON
Volume 106, pp. 103-108 October-December 2008
Mucor guilliermondii (Mucorales): a rare species
found in herbivore dung from Neotropics
ANDRE LUIZ CABRAL M. DE A. SANTIAGO’,
Marta A. Q. CAVALCANTI’ & SANDRA FARTO BOTELHO TRUFEM?
andrelcabral@msn.com, xiliamac@terra.com.br, strufem@cy.com.br
‘Programa de Pos-graduacao em Biologia de Fungos. Universidade Federal de
Pernambuco. Av. Prof. Nelson Chaves, s/n, 50670-420, Recife, PE, Brasil
?Universidade Sao Marcus
Rua Antonio Gomes Ferreira 89, 04257-100, Sao Paulo, SP. Brasil
Abstract — Mucor guilliermondii was isolated for the first time in Neotropics, from
tapir (Tapirus terrestris) dung collected at the Reserva Ecoldgica de Dois Irmaos
(8°7°30” S and 34°52°30” W), located in Recife, State of Pernambuco, Northeast Brazil.
An identification key for species of Mucor studied in Brazil is provided.
Key words — Zygomycetes, Mucoraceae, taxonomy
Introduction
Mucor includes about 80 cosmopolitan species and is the most representative
and studied genus of Mucoraceae (Domsch et al. 1995). This genus belongs
to the subphylum Mucoromycotina (Hibbett et al. 2007), order Mucorales
family Mucoraceae (Benny 2005) and contains many coprophilous and
noncoprophilous species (Krug et al. 2004).
Mucor guilliermondii was described by Nadson & Philippov (1925) from
Periplaneta americana dung in Russia and re-described by Schipper (1978).
There are few reports of isolation from soil in Malaysia (IMI 350568) and from
humus horizon in Belgium (MUCL 38359). This species is differentiated from
the other Mucor species by the production of elongated-conical columellae
with extremely small or absent collar and ellipsoidal, cylindrical to reniform
sporangiospores, generally with a droplet at each end (Schipper 1978).
The aim of this work was to describe M. guilliermondii isolated from tapir dung
in Recife, PE, and to provide an identification key for Mucor species studied in
Brazil.
104 ... Santiago, Cavalcanti & Trufem
Materials and methods
The samples of tapir (Tapirus terrestris) dung were collected at Reserva Ecologica
de Dois Irmaos (8°7’30”S and 34°52’30”W), which includes a Zoological Park
and is located in Recife, State of Pernambuco, Northeastern Brazil. The area
contains 387 ha and is an Atlantic Forest Ecological Reserve.
The samples of dung were collected with a sterilized spatula, placed in plastic
bags, taken to the laboratory and incubated in moist chamber at 28°C + 2°C
for 7 days. |
Fragments of mycelium were removed directly from the samples at the
stereomicroscope and transferred to the Petri dishes with potato dextrose
agar (PDA) (Lacaz et al. 2002) plus chloramphenicol (80 mg/L). Identification
was based on macroscopic (color, aspect and diameter of the colonies) and
microscopic (microstructures) characters according to Nadson & Philippov
(1925) and Schipper (1978).
Taxonomy
Mucor guilliermondii Nadson & Philippov,
Rev. Gén. Bot. 37: 450 (1925) Fig. 1 a-d
Colony with limited growth after 6 days at 26°C in PDA, 6.5 cm diam and 3
mm height. Colony white with cottony aspect, irregular edges and yellowish
reverse. Aroma lightly sweet. SPORANGIOPHORES not ramified, 2.5—6 um diam,
erect, some bending, soon collapsing. SPORANGIA hyaline with vitreous aspect,
yellow, globose and subglobose to lightly depressed, 14.5-52.5 um diam with
smooth, transparent and deliquescent wall. COLUMELLAE elongated-conical,
rare subglobose and ellipsoidal with lightly incrusted wall, 15-30 x 5.5-14.5 um,
majority with 15-20 x 5.0-8.5 um, small or absent collar. SPORANGIOSPORES
hyaline, smooth walled, 3.5-8 x 2.0-4.5 um, ellipsoid and cylindrical to
reniform, some irregular, with a droplet at each end. Some cells may contain
more than two droplets. CHLAMYDOSPORES globose, 12.5-23 tum, subglobose,
cylindrical and doliiform, 10-22.5 x 8-22 um. ZyGosPorEs not observed.
Hasirat: Tapir dung (Tapirus terrestris).
GEOGRAPHIC DISTRIBUTION: Russia (CBS 174.27), Malaysia (IMI 350568),
Belgium (MUCL 38359). This is the first report in Neotropics.
SAMPLE INFORMATION: Brazil, Pernambuco, Recife, Reserva Ecologica de Dois Irmaos,
May/06, A. L. C. M. A. Santiago (URM5848).
REMARKS: The characteristics of the strains of M. guilliermondii reported here
show a close similarity with the descriptions of Nadson & Philippov (1925)
and Schipper (1978). However, differences on colony color and diameter of
sporangiophores were observed. Nadson & Philippov (1925) reported wider
Mucor guilliermondii in Brazil... 105
Fic. 1 Mucor guilliermondii
A) Sporangiophore; B) Sporangium; C) Columella; D) Sporangiospores.
Scale bars: A = 40 um; B = 20 um; C = 4 um; D = 5 um.
sporangiophores (7-10 tm diam) in relation to the ones described by Schipper
(1978; up to 7 um diam) and those described in this manuscript (2.5-6 um
diam). The colony color may vary from white (Nadson & Philippov 1925)
106 ... Santiago, Cavalcanti & Trufem
to pale gull gray (Schipper 1978). Although sporangiospores usually show a
droplet at each end, according to Nadson & Philippov (1925) the number of
droplets may vary (3, 4 or more).
Mucor guilliermondii was treated as a synonym of M. subtilissimus Oudem. by
Zycha (1969). According to Schipper (1978) ‘the almost obclavate columellae in
M. guilliermondii are hard to reconcile with the columellae of M. subtilissimus,
which were described by Oudemans (1898) as being globose. The globose
columellae on M. subtilissimus was also observed by Alves et al. (2002) who
pointed out that M. subtilissimus sporangiophores present a constriction next
to young sporangia or swelling 50 um below the columellae, which are not
visualized in M. guilliermondii.
This manuscript reports for the first time the occurrence of Mucor guilliermondii
in Neotropics, contributing with the knowledge of Mucorales distribution.
In Brazil, other species of Mucor such as M. circinelloides Tiegh., M. fuscus
Bainier, M. genevensis Lendn., M. hiemalis Wehmer, M. mousanensis Baijal &
B.S. Mehrotra, M. mucedo L., M. piriformis A. Fisch., M. plumbeus Bonord., ©
M. racemosus Fresen., M. subtilissimus, and M. variosporus Schipper have been
isolated from herbivore dung (Batista et al. 1961a, b, c; Trufem 1984, Trufem &
Viriato 1985, Alves et al. 2002).
Identification key for species of Mucor from herbivore dung in Brazil
L)Zygospores presentin monosporic cultures’... 22... ee M. genevensis
l= Zygospotes absent in monosporic Cultures. ae ov.tad ee eee ee ys
2. Mycelia delicate, sporangiospores with one or more droplets at each end.......... 2
2. Mycelia vigorous, sporangiospores without droplets at the ends.................. 4
3. Columellae globose, sporangiophores with a constriction next to young
sporangia or swelling 50 um below the columellae............ M. subtilissimus
3. Columellae elongated-conical, sporangiophores not as above..... M. guilliermondii
A=Sporangiaswith persistent: wallace ches. 7.) aye ei a eee eer 5
4ySporangia with deuquescent Wall % 22004 ges. a/s ts See ye oo ys eat ete eee 6
5. Columellae globose, obovoid; sporangiospores variable in shape and size
ET Ng Ee EIR Ee RTF SOE cg A, M. variosporus
5. Columellae elongated-conical or ellipsoidal; sporangiospores ellipsoidal,
repular in SiZe7 9 We eee ce a ra ee re M. mousanensis
6 sporangiaioiien more.than. 150g diane wee eee een te ee ir,
6. Sporangia‘otten lessithiand 50 nt diane ter eae Se ee tg 8
7. Sporangiospores ellipsoidal up to 10 um in length; rapid growth on cherry agar
Or acid, DECEWOLL A0al le. Aik ee ee ee eee LT, M. piriformis
7. Sporangiospores cylindrical-ellipsoidal, often over 10 um in length; restricted
growth on cherry agar or acid beerwort agar...................--- M. mucedo
Mucor guilliermondii in Brazil ... 107
8. Sporangiophores unbranched or weakly sympodially branched ................... 9
8. Sporangiophores repeatedly sympodially branched.....................000000- 10
9. Columellae globose; sporangiospores ellipsoidal, fusiform ..... M. hiemalis f. luteus
9. Columellae subglobose with a truncate base, some subglobose; sporangiospores
ellipsoidal, sometimes flattened at one side ............. M. hiemalis f. hiemalis
10. Chlamydospores abundant in sporangiophores........ M. racemosus f. chibinensis
mim latiyCOspores tate imysporangiGpMoresrem tam ty ane Put. Be sith oh, ihe ee iat
11. Sporangiospores globose and globose to subglobose, rare irregular............. 12
11. Sporangiospores subglobose and ellipsoidal, some globose.................... 13
12. Sporangiophores branching sympodially and monopodially; columellae
Pile Lat Dipl Oj CCLLO nim Wares Aare Berwin caren ds) ht ates Su 8 ays M. plumbeus
12. Sporangiophores branching sympodially; columellae rarely with
DIOJOCHIONS serey use, 2 A Ot. Ae a ete meee ao FUE 38 M. fuscus
Meme@olonics whiteatifirstylatereray.auz6@ in PDAL a. ee ee 14
Beeeolunics white at first, later brown at 26°C in PDA 2 eee. fe.. cee ibs
14. Sporangiospores subglobose, some globose 7 um diam
as Tn Wc hie aia atch een dee oly alien cette Pers Mee eW M. circinelloides f. janssenii
14. Sporangiospores ellipsoidal, 5-11.2 x 2.5-6.2 um
eg ees Le CER Ne Oe es ae M. circinelloides f. griseocyanus
15. Columellae obovoid to ellipsoidal, globose in the smaller sporangia
4s; gaa bs Rael AUN a ee eae hae? fo Rae M. circinelloides f. circinelloides
Bemomellaeplobosemrwyte wake oes a eee M. circinelloides f. lusitanicus
Acknowledgements
The authors thank Dr José Luiz Bezerra and Dr Hsiao-Man Ho for article review, Felipe
Vasconcelos Reis for English review, Coordenacao de Aperfeicoamento de Pessoal
de Nivel Superior (CAPES) for a PhD Scholarship provided to the first author, and
Conselho Nacional de Desenvolvimento Cientifico e Tecnolégico (CNPq) for providing
a research grant to the second author.
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MYCOTAXON
Volume 106, pp. 109-126 October-December 2008
The genus Stereum in Israe!
DANIEL TuRA"*, IVAN V. ZMITROVICH?,
SOLOMON P. WASSER?? & EVIATAR NEVO’
*turadaniel21@yahoo.com
‘Department of Evolutionary and Environmental Biology, Institute of Evolution
Faculty of Science and Science Education, University of Haifa
Mt. Carmel, Haifa 31905, Israel
*Lab. of the Systematics and Geography of Fungi, Komarov Botanical Institute
Russian Academy of Sciences, Prof. Popov Str.2, St. Petersburg, 197376, Russia
°N.G. Kholodny Institute of Botany, National Academy of Sciences of Ukraine
Tereshchenkivska Str. 2, Kiev 01001, Ukraine
Abstract — A study of Israeli Stereum species diversity resulted in four species: Stereum
gausapatum, S. hirsutum, S. sanguinolentum, and S. subpileatum. A new form, S.
hirsutum f. lobulatum, is described from the species complex, and S. gausapatum and S.
sanguinolentum are recorded as new species for Israeli mycobiota. Taxa are described
and microstructural features are illustrated based on specimens collected from northern
Israeli forests. Notes on host preference, morphological variability, distribution, and
taxonomic relationships are provided. A key to Israeli Stereum species is given.
Key words—Mediterranean, stereoid fungi, polymorphism, taxonomy
Introduction
Its xerophilic habit makes the genus Stereum (Stereaceae, Russulales) a highly
characteristic indicator for arid mycotas. This genus includes white-rot fungi
inhabiting fallen or standing branches, trunks, and stumps representing a
broadly diverse selection of broadleaved and coniferous trees. Their economic
importance lies in their wood-destroying properties and carbon recycling of
arid forest ecosystems (Overholts 1939, Woon & Jung 1999).
The stereoid morphotype covers annual to perennial, tough-leathery
substipitate to resupinate fruitbodies with a more or less even hymenophore.
Generically, the Stereum pileal covers are trichodermoid, often transforming
to a fomitoid crust. The hymenium is variably colored, occasionally staining
yellow-orange or red when injured. Cross-sections of the cream-colored context
11.0 .... Tora, &al:
reveal thin dark reddish deposits between the tomentum and fruitbody core.
Russuloid pseudocystidia with refractive contents and weakly amyloid spores
are characteristic at the microscopical level. The hyphal system is variously
described as dimitic or monomitic, but most species have a pseudodimitic
hyphal system dominated by thick-walled hyphae with living protoplasts. All
hyphae are densely packed so that the fruitbodies are rather hard.
The concept of Stereum, adopted by Persoon (1794) and Fries (1838), was
long used as an infrageneric division of Thelephora Ehrh. On the other hand, the
concept of this union was rather broad. After generic separation and intensive
microscopical studies, the concept of Stereum became increasingly narrow.
Lentz (1955), Donk (1957), Boidin (1958a, b, 1959a, b, 1960), Pouzar (1959),
Parmasto (1968), Welden (1971), and Chamuris (1988) segregated Stereum
into smaller genera, thereby making the remaining genus a more homogeneous
group. However, some of these segregate genera (e.g., Haematostereum Pouzar,
Xylobolus P. Karst. emend. Boidin) have been “merged” into the core Stereum
group, creating additional taxonomical problems. Some Xylobolus characteristics
— such as amyloid binucleate basidiospores, hymenial acanthophysoid -
elements, and homothallism — are also basic generic characters for Stereum,
suggesting that segregation of the genus has been premature.
Species concepts within Stereum are complicated by a lack of stable
character patterns. Basidiocarp shape, type of tomentum, and presence of
acanthohyphidia and conducting hyphae (Jiilich & Stalpers 1980).
Modern molecular investigations place Stereum in the russuloid clade,
which fully corresponds with microscopic evidence. Characteristics such
as amyloid reactivity of the spore wall and the presence of gloeoplerous and
lactiferous hyphae in most Stereum species are considered homologous with
other russuloid taxa (Larsson & Larsson 2003, Larsson 2007).
The genus Stereum is distributed worldwide, with many widespread species
following the distribution of their hosts from the boreal zone of Europe and
North America into the arid areas of Africa, Asia, Australia, and the Middle
East (Binyamini 1982, 1984; Breitenbach & Kranzlin 1986; Chamuris 1988;
Ginns & Lefebvre 1993).
The history of Stereum research in Israel began with Binyamini (1982), who,
in his fourth paper on Israeli lignicolous Aphyllophorales, recorded Stereum
hirsutum and Xylobolus subpileatus (later described in Binyamini 1984) as
species new to the area. Stereum hirsutum, considered a complex by many
authors due to its great morphological variability, occurs on various hardwood
substrata, shows a high ecological tolerance, and is also widespread in northern
Israeli broadleaf forests. In contrast, Stereum (Xylobolus) subpileatum has been
found only once growing on Quercus calliprinos in the Samarian forests. We
have also found Stereum gausapatum inhabiting Q. calliprinos stumps in the
Stereum in Israel... 111
Israeli mountain forests of the Golan Heights. This species follows its host,
usually oak, in Europe, Asia, Australia, and North America (Breitenbach &
Kranzlin 1986). We present the first reports of Stereum gausapatum as well as
the conifer-inhabiting S. sanguinolentum (found growing on fallen pine wood
in the Golan Heights) for Israel below.
Material and methods
Fruitbody morphology was examined under light/dark
field using a Carl Zeiss Axiostar 1122-100 microscope.
Measurements were made using an immersion objective;
30 spores were measured for each specimen. The highest
and lowest 5% spore measurements (given in parentheses)
were excluded from each size variation range as outliers.
Tissues were mounted in 10% potassium hydroxide (KOH)
aqueous solution and Melzer’s reagent for microscopical
analyses. The material used in our analyses can be found
in the herbarium of the Institute of Evolution, University
of Haifa (Haifa, Israel, HAI). A map (Fic. 1) shows the
Israeli distributions of each newly recorded species were
distributed (Feinbrun-Dothan & Danin 1998).
Fic. 1. Accepted abbreviations of nature regions of Israel: AP - Akko Plain; AV - Arava Valley; BS
- Beit Shean Valley; CC - Carmel Coast; CG - Coast Galilee; CM - Carmel Mount; CN - Central
Negev; DS - Dead Sea Area; EP - Esdraelon (Yizreel) Plain; GH - Golan Heights; GM - Gilboa
Mount; HE - Hermon Mount; HP - Hula Plain; JD — Judean Desert; JM — Judean Mts.; LG — Lower
Galilee; LJ - Lower Jordan Valley; NN - Northern Negev; PP - Philistean Plain; SA —- Samaria; SH
- Shefela; SN - South Negev; SP - Sharon Plain; UG - Upper Galilee; UJ - Upper Jordan Valley;
WN - Western Negev.
Taxonomic descriptions
Stereum gausapatum (Fr.) Fr.. Hymenomyc. Eur.: 638, 1874. FIGS. 2-3
= Thelephora gausapata Fr., Elench. Fung. 1: 171, 1828.
BASIDIOCARPS annual, coriaceous, sessile to resupinate, normally effused-
reflexed, 3-15 cm in extent, on vertical substratum found as imbricate
dimidiate pilei merging into board-like patches. ABHYMENIAL SURFACE
tomentose to strigose-hirsute; slightly zonate, rust-brownish with white-cream,
and thin inflexed undulating margin. HYMENIAL SURFACE even to somewhat
tuberculate, cracking, brittle and pale-brown when fresh, but gray-brown when
dry; when injured, fresh basidiome bruising red. CONTExT thin, homogeneous,
0.5-1 mm. thick, cream-colored, in dry specimens without an evident deposit
between tomentum and context.
YQ. eure)
Fic. 2. Stereum gausapatum growing on Quercus calliprinos stump in GH, Israel.
Scale bar = 1 cm (Photo V.F. Malysheva).
Fic. 3. Microscopic structures of Stereum gausapatum:
a — hyphae; b - hymenial fragment showing slenderly thick-walled pseudocystidia
among cylindrical basidioles, basidia, and basidiospores.
Scale bar = 10 um.
Stereum in Israel... 113
thus individual hyphae hardly seen, while contextual hyphae are somewhat
loose. Generative hyphae septate, thin- to slightly thick-walled, 2-4 um diam,
branched and hyaline. Pseudoskeletal hyphae thick-walled, hyaline to yellowish
with oily contents inside, slenderly branched, 4-8 (mostly 4-5) um diam.
Conducting hyphae similar, but without septation, with chrysescent contents.
CysTIDIA not seen, but cystidia-like hyphal ends (pseudocystidia) numerous,
cylindrical to subcylindrical, thin- to slightly thick-walled, 2-7 um wide (with
basal inside walls 1-1.5 um thick), but apically thin-walled, with brownish oily
contents. BASIDIOLES cylindrical, thin-walled, 2-3 um diam, projecting up to
10 um above hymenium. Basrp1a4 narrowly clavate to clavate, 43-55 x 6-10 um,
4-spored, without a basal clamp. Basip1osporss ellipsoid-cylindrical, hyaline,
smooth, thin-walled, (6-)6.5-7.9 x 3-3.8 um, amyloid.
: Found growing on Quercus calliprinos in the North of Israel (Fic. 4).
|
HyPHAL SYSTEM pseudodimitic. Subhymenial hyphae strongly interwoven,
GENERAL DISTRIBUTION AND HABITAT. Stereum gausapatum
occurs mostly in oak zones on Quercus, Castanea, and Carpinus *| “ ;
species. Basidiomata develop on dead stems, rotten stumps,
logs, or trunks. Associated with a white rot (Ginns & Lefebvre ..
1993). Its general distribution according to our knowledge
includes: Europe (Austria, Belgium, Denmark, Estonia,
France, Germany, Georgia, Greece, Norway, Portugal, Russia, ©
Spain, Sweden, Ukraine); Mippie East (Israel, Turkey), Asia
(Azerbaijan, Japan, South Korea); NoRTH AMERICA (Mexico, « ; 5
USA); SouTH AMERICA (Panama).
Fic. 4. Distribution of Stereum gausapatum in Israel.
Note. Stereum gausapatum was considered one of the “bleeders’, along
with S. sanguinolentum and S. rugosum Pers. and therefore placed into
Haematostereum by Pouzar (1959). The bleeding reaction in fresh specimens
appears upon hymenial surface injury and is caused by inner-colored exudates
of the numerous pseudocystidia present in the hymenium of these species
(Fig. 17). Due to this particular fact, S. gausapatum can be confused with S.
rugosum, which when bruised also stains red and in addition shares the same
habitat, however differs by its stratified perennial hymenium, larger spores, and
presence of acanthohyphidia. Other species sharing a common habitat with S.
gausapatum, are S. hirsutum and S. ostrea (Blume & T. Nees) Fr. In the field, S.
gausapatum can be distinguished from S. ostrea by densely hirsute and thicker
pilei (Overholts 1939), but, microscopically, S. gausapatum does not possess
acanthohyphidia. In Israel, S. gausapatum may be confused with specimens
belonging to S. hirsutum-complex because they share the same habitat, have
similar spore sizes, and, in some specimens, turn red when injured during their
114... Tura &al.
active growth phase. However, S. hirsutum has thick-walled pseudocystidia and
usually acute hyphidia, while the pseudocystidia in S. gausapatum have rather
thin walls: less than 1.5 um in diam. (Chamuris 1985, 1988). According to Jiilich
& Stalpers (1980), within S. gausapatum, the dimension of the spores given by
European authors is always larger than those given by American authors.
SPECIMENS EXAMINED: ISRAEL. GH, Massada Forest, Quercus calliprinos forest, on
Quercus stump, 07.03.2007, leg. & det. D. Tura, I. Zmitrovich et V. Malysheva (HAI
0114).
Stereum hirsutum (Willd.) Pers., Observ. Mycol. 2: 90, 1800 (‘1799’). Fics. 5-9
= Thelephora hirsuta Willd., Fl. Berol. Prodr.: 397, 1787.
BASIDIOCARPS annual to perennial, usually sessile to effused-reflexed with
narrow to broadly prostrate base or totally resupinate and several centimeters
in extent. Pilei semicircular, dimidiate or flabelliform to irregular, often
umbonate, up to 70 x 50 x 1-2 mm; membranous-elastic when fresh and
ceraceo-coriaceous when dry. When broadly attached, in most cases with
confluent pilei, the single pilei have an imbricate arrangement. ABHYMENIAL |
SURFACE hirsute to hispid, yellow-ochraceous or grayish-white, in older
specimens often covered by green algae or with a reddish-brown cutis exposed
towards the attachment zone, more or less zonate. When zonate, usually with
narrow dark bands alternating with wider and lighter colored ones (zonation in
some specimens may be more prominent towards margin, where cutis is visible
as a narrow glabrous band). Margin wavy, sometimes radiately plicate, thinning
outwards, usually whitish-cream to gray. HYMENIAL SURFACE smooth to slightly
tuberculate, often with radial concentric ridges; light-yellow to orange-grayish,
when old hymenium turns gray, in some specimens growth may continue on
the same-aged basidiocarp; thus, sometimes marginal hymenium appear to be
thicker and lighter in color. CONTEXT thin, whitish-cream; in cross section,
with a very thin red-brownish deposit between tomentum and context, barely
observable in some specimens.
HyPHAL SYSTEM pseudodimitic. Generative hyphae simple septate, thin-
walled, mostly 2—3um diam, branched and hyaline; pseudoskeletal hyphae
thick-walled, rarely septate, 4-8 um diam, unbranched to slightly branched,
with oil-drops and pale yellowish contents in KOH. Cystip1a not seen, but
cystidia-like hyphal ends (pseudocystidia) present. PsEUDOCYSTIDIA abundant,
cylindrical to subcylindrical, sometimes medially inflated, thick-walled, except
an apical part with basal wall 3-4 um thick. HypHip1a acuminate, mostly
cylindrical, thin- to slightly thick-walled, 2-3 um in diam. Basip1a slenderly-
clavate, 25-35 x 3.5-4 um, 4-spored, without a basal clamp. BASIDIOSPORES
ellipsoid-cylindrical, hyaline, thin-walled, even, often with internal oil-drops,
4—6.5(-7) x 3-3.5 um, amyloid.
Stereum in Israel... 115
Fig. 5. Microscopic structures of Stereum hirsutum:
a — hyphae; b —- thick-walled pseudocystidia; c - hymenial fragment showing pseudocystidia
among acute-cylindrical hyphidia, basidia, and basidiospores.
Scale bar = 10 um.
GENERAL DISTRIBUTION AND HABITAT. The species within the Stereum
hirsutum-complex are generally known to grow on bark and wood of trunks,
twigs, or stumps of a wide range of dead or living deciduous trees. According
to present-day knowledge, the members of this complex are widespread and
common in several countries: EUROPE (Austria, Croatia, Denmark, Finland,
France, Germany, Greece, Iceland, Italy, Macedonia, the Netherlands, Norway,
Poland, Portugal, Romania, Russia, Spain, Sweden, UK, Ukraine); MIDDLE
East (Israel, Turkey); East Asia (Mongolia, Japan, South Korea); NorTH
AMERICA (Canada, USA); SouTH AMERICA (Colombia, Costa Rica, Dominican
Republic, Venezuela); SOUTHERN HEMISPHERE (Australia, New Zealand);
AFRICA (Morocco, Tanzania); GREATER ANTILLES (Jamaica).
Note. Some authors treat Stereum hirsutum as a complex that shares a common
or similar inner structure but exhibits different morphological forms; other
authors find “reliable” taxonomic features to split this complex into separate
species. However, some specimens, depending on biotic and abiotic factors of
their habitat in various geographic biomes, show some stable morphotypes.
During its active growth, hymenium in some fruitbodies turns yellow-orange
to red when bruised; thus, in the field, they might be confused with S. rugosum
and S. gausapatum. Therefore, careful morphological examination is needed.
Because of its multiplicity of forms, according to Breitenbach & Kranzlin
(1986), old specimens of S. hirsutum can be confused with S. ochraceoflavum
(Schwein.) Fr. Although both species share the same substrata, S. ochraceoflavum
116... Tura & al.
preferentially colonizes twigs and clippings, while S. hirsutum colonizes trunks,
crevices, and thick branches and thus apparently produces thicker fruitbodies.
Chamuris (1988) cites the following important stable characters for the
North American S. hirsutum (all also found in Israeli examined specimens):
the presence of cutis, broad substratum preference, mostly acuminate hyphidia,
thick-walled pseudocystidia with basal wall thickness = 2 um wide, and
conducting hyphae with yellow contents that turn brownish-orange in Melzer’s
reagent.
Stereum hirsutum is one of the most common hardwood-inhabiting species
in Israel. The variable morphologies exhibited by the species in northern Israeli
forests are compared in TABLE 1.
TABLE 1. Fruitbody variability in Stereum hirsutum-complex in Israel
MORPHOTYPES BASIC CHARACTERISTICS
Strongly fruitbodies strongly plicate, imbricate, narrowly to sessile attached;
plicate, actively abhymenial surface tomentose-strigose ocher-pale brownish, towards attachment
growing zone covered by green algae or with a glabrous reddish brown cutis;
type hymenium even, with radially concentric ridges, yellowish-orange.
Note: The yellow to reddish bruising hymenium (sometimes, when fresh)
HAI 0141 suggests an actively growing basidiocarp; usually, in such cases, the abhymenial
(Fie. 6) and hymenial surfaces show lighter colors compared to aged fruitbodies (Fig. 7).
Aged fruitbodies narrowly attached to sessile, dimidiate, flabelliform;
fruitbody abhymenial surface white-gray, zonate, tomentose-strigose, with cream- colored
type margin and often glabrous, blackish-brown cutis towards attachment zone;
hymenial surface yellow-orange to gray, smooth.
Note: the hard, dry fruitbodies, the glabrous, brown-blackish visible cutis,
HAT 0119 often covered by green algae and the presence of grayish color on hymenial and
(Fic. 7) abhymenial surfaces resemble the main features of aged fruitbodies.
lneven fruitbodies effused-reflexed, narrowly attached, confluent;
hymenophore abhymenial surface zonate, mated-strigose, grayish;
type hymenial surface even-tuberculate beset with minute, irregular, lighter colored
folds, which actually resemble small, rising individual pileoli with light-colored
patches on a grayish background.
HAT 0138 Note: only two samples with this type of deviant hymenial surface, were found
(Fie. 8) in Israel in different localities.
fruitbodies imbricate, broadly to narrowly attached in appearance, 90 x 17 x 1 mm
Tomentose thick, confluent; an individual pilei with its own attachment point;
abhymenial abhymenial surface tomentose, tomentum entirely covering the upper surface -
surface including short stipe, zonate;
ee hymenial surface even, light yellowish-orange to grayish towards attachment
point.
HAI 0137 Note: the soft tomentum covering the entire upper surface and the warm light
(Fic. 9) colors make this specimen highly distinguishable from the other samples of
Stereum that we found.
Stereum in Israel... 117
Fic. 6. Stereum hirsutum fruitbody: strongly plicate, actively growing type
(explanation in TABLE 1). Scale bar = 1 cm.
Fic. 7. Stereum hirsutum fruitbody: aged fruitbody type
(explanation in TABLE 1). Scale bar = 1 cm.
118 ... Tura & al.
Fia. 8. Stereum hirsutum fruitbody: uneven hymenophore type(explanation in TABLE 1).
Scale bar = 1 cm.
€ .
Fig. 9. Stereum hirsutum fruitbody: tomentose abhymenial surface type (explanation in TABLE 1).
\ WS SSS
S)
Scale bar = 1 cm.
Chamuris (1988) suggests that S. subtomentosum Pouzar gradually merges
into S. hirsutum. Indeed, in boreal areas, specimens characterized by applanate
symmetrical pilei with dull-colored hymenium can be found, but the presence
or absence of a strigose abhymenial surface greatly complicates species
attribution. However, as Israeli materials, despite their variability, do not
include S. subtomentosum-like morphotypes, this taxon may be considered
absent in Israel.
The S. hirsutum species complex is widely distributed in North Israel (Fic.
10).
Stereum in Israel ... 119
Fic. 10. Distribution of Stereum hirsutum-complex in Israel.
Many Israeli specimens of S. hirsutum exhibit longitudinally
expanded lobate processes, a very interesting feature that
is seen elsewhere only in S. ostrea (incl. S. lobatum) and S.
insignitum Queél. Its strigose pilei with a black line under the
tomentum and the absence of acanthohyphidia, confirms the
identification of the Israeli material as S. hirsutum, yielding
an ecotype parallel to S. ostrea and S. insignitum. As this
pattern is very distinguishable, we propose a new form for this
polymorphic species:
Stereum hirsutum f. lobulatum Tura, Zmitr. & Wasser, f. nov. FIGs. 11-12
MycosBaANnk MB 506905
Basidiomata stereoidea, perennia, basi angustata, lobulata, elongata, ad marginem
undulata et confluenta. Superficio strigosa, zonata et undulata, basi cinerascens, ad
marginem ochraceo-flava. Hymenium laeve, sed radiately costatum, basi cineraceo-
flavum, ad marginem subvitellinum. Structura microscopica quod in typus. Sporae 5.5-7.5
x 2.5-3.0 pm.
Ho.otype: ISRAEL. UG, Savsufa Forest, Quercus calliprinos forest, on oak stump,
11.02.2008, leg. D. Tura, det. D. Tura et I. Zmitrovich (HAI 0193).
BASIDIOCARPS perennial, usually sessile with narrow base, 1-2.5 cm wide and
1.5-3.5 cm long; pilei flabelliform to irregular, regularly lobate, confluent both
in basal and marginal parts. ABHYMENIAL SURFACE hirsute to hispid, grayish
at the base and yellow-ochraceous at margin, regularly zonate, radiately plicate.
HYMENIAL SURFACE basically smooth but with radial concentric ridges along
fruitbody growth axes, light-yellow to orange-grayish. CONTEXT thin, pale
cream; in cross-section with a very thin cinnamomeous deposit between
tomentum and context.
The microstructure is the same as in all other specimens of S. hirsutum
studied; acanthohyphidia absent. Spores 5.5-7.5 x 2.5-3.0 um.
Note. Such a specific lobulate and subfasciate habit has not been described for
Stereum hirsutum, at least not in well-known monographs (Eriksson et al. 1984,
Davydkina 1980, Chamuris 1988, Duhem 2007). Chamuris (1988), who noted
possibly undulate or plicate fruitbody forms, did not mention deeply crenate
lobules in his monograph.
All lobulate forms of this fungus were found in subxerophilic mountain
oak forests (Upper Galilee, Golan Heights). In such extreme conditions, the
fungal morphogenetic spectra are enriched by strange “marginal” forms. Some
of these (e.g., Byssomerulius corium var. halieensis Zmitr. et al., Peniophora
120 ... Tura & al.
Ss
Fig. 11. Stereum hirsutum f. lobulatum fruitbodies variability.
Cl Sle Fae
Fig. 12. A large-scale view of fruitbody cluster of Stereum hirsutum f. lobulatum.
Se
Stereum in Israel... 121
quercina f. merulioides Tura et al.) have been described earlier so as to highlight
the variability range of these species (Zmitrovich et al. 2006, Tura et al. 2007).
These formations are very important for an internal differentiation of the
species because over time reproductive barriers may occur within such ecotypic
subpopulations that may lead to eventual speciation.
SPECIMENS EXAMINED: ISRAEL. UG, Mt. Meron National Park, on Pinus?, 27.12.2000,
leg. S. Reshetnikov, det. S. P. Wasser (HAI 0115); ?, Harshad Harbaim, on branch of dead
oak, 1.02.2001, leg. S. Reshetnikov, det. S. P. Wasser (HAI 0116); ?, Harshad Harbaim,
on Quercus stump, 24.02.2001, leg. I. Tovbin, det. S.P. Wasser (HAI 0117); UG, Salsufer
Forest, on Q. calliprinos wood, 19.11.2006, leg. D. Tura, det. D. Tura, I. Zmitrovich &
V. Malysheva (HAI 0118); UG, Bar'am Forest, Q. calliprinos forest, on Quercus stump,
16.12.2006, leg. D. Tura, det. D. Tura, I. Zmitrovich et V. Malysheva (HAI 0119); GH,
Massada Forest, Q. calliprinos forest, on broadleaf wood, 26.11.2006, leg. D. Tura,
det. D. Tura, I. Zmitrovich & V. Malysheva (HAI 0120); LG, Oshaya Forest, mixed
Pinus and Quercus wood, on Quercus stump, 16.12.2006, leg. Dalurasdet. De luran.
Zmitrovich & V. Malysheva (HAI 0121); UG, Nahal Keziv, mixed hardwood forest, on
Quercus stand, 20.12.2006, leg. D. Tura, det. D. Tura, I. Zmitrovich & V. Malysheva (HAI
0122); MC, Muhraqua, Q. calliprinos forest, on Quercus stump, 24.12.2006, D. Tura, I.
Zmitrovich et V. Malysheva (HAI 0123); GH, Qazrin, mixed forest, on Cupressus stump,
09.01.2007, leg. D. Tura, det. D. Tura, I. Zmitrovich & V. Malysheva (HAI 0124); GH,
Massada Forest, Q. calliprinos forest, on broadleaf wood, 09.01.2007, leg. D. Tura, det. D.
Tura, I. Zmitrovich & V. Malysheva (HAI 0125); MC, Mt. Carmel Natural Park, mixed
Quercus and Pinus forest, on Quercus branches, 12.01.2007, leg. S. Zohar, det. D. Tura,
I. Zmitrovich & V. Malysheva (HAI 0126); UG, Even Menahem, Quercus forest, on
hardwood stump, 22.01.2007, leg. Y. Ur, det. D. Tura, I. Zmitrovich & V. Malysheva (HAI
0127); UG, Hanita Forest, Q. calliprinos forest, on branches of Q. calliprinos, 31.01.2007,
leg. Y. Ur, det. D. Tura, I. Zmitrovich & V. Malysheva (HAI 0128); LG, Lavi Forest, mixed
Pinus and Quercus forest, on oak twig, 04.02.2007, leg. Y. Ur, det. D. Tura, I. Zmitrovich
& V. Malysheva (HAI 0129); UG, Kiryat Shmona, Eucalyptus wood, on hardwood twigs,
11.02.2007, leg. Y. Ur, det. D. Tura, I. Zmitrovich & V. Malysheva (HAI 0130); SA, Reihan
Forest, mixed Pinus and Quercus wood, on hardwood twigs, 19.02.2007, leg. Y. Ur, det.
D. Tura, I. Zmitrovich & V. Malysheva (HAI 0131); UG, Kiryat Shmona, Eucalyptus
wood, on hardwood stump, 20.02.2007, leg. D. Tura, det. D. Tura, I. Zmitrovich & V.
Malysheva (HAI 0132); UG, Banias Park, mixed broadleaf trees, 20.02.2007, leg. Yair Ur,
det. D. Tura, I. Zmitrovich et V. Malysheva (HAI 0133); UG, Sde Eliezer, on Eucalyptus
tree, 21.02.2007, leg. Y. Ur, det. D. Tura, I. Zmitrovich & V. Malysheva (HAI 0134); GH,
Massada Forest, Q. calliprinos forest, on oak stump, 07.03.2007, leg. D. Tura, det. D. Tura,
I. Zmitrovich & V. Malysheva (HAI 0135); UG, Kiryat Shmona, Eucalyptus wood, on
hardwood stump, 07.03.2007, leg. D. Tura, det. D. Tura, I. Zmitrovich & V. Malysheva
(HAI 0136); LG, near Kabul, mixed hardwood forest, on Eucalyptus stump, 07.03.2007,
leg. D. Tura, det. D. Tura, I. Zmitrovich et V. Malysheva (HAI 0137); UG, Mt. Meron,
Q. calliprinos forest, on Quercus stump, 09.03.2007, leg. & det. D. Tura, I. Zmitrovich &
V. Malysheva (HAI 0138); UG, Bar’am Forest, Q. calliprinos forest, on Quercus stump,
09.03.2007, leg. D. Tura, det. D. Tura, I. Zmitrovich & V. Malysheva (HAI 0139); SP,
Ben Shemen Forest, mixed broadleaf trees, on ?, 16.03.2007, leg. Y. Ur, det. D. Tura, I.
Zmitrovich & V. Malysheva (HAI 0140); UG, Canyon Agam, Quercus forest, on Quercus
stump, 17.03.2007, leg. Zohar S., det. D. Tura, I. Zmitrovich & V. Malysheva (HAI 0141);
GH, Hanita Forest, Q. calliprinos forest, on branches of Q. calliprinos, 20.03.2007, leg. Y.
Ur, det. D. Tura, I. Zmitrovich & V. Malysheva (HAI 0142).
1220 ae ldrascal
Fic. 13. Dry fruitbodies of Stereum sanguinolentum on fallen pine wood.
Scale bar = 1 cm.
Stereum sanguinolentum (Alb. & Schwein.) Fr., Epicr. Syst. Mycol.: 549, 1838.
FIGS. 13-15
= Thelephora sanguinolenta Alb. & Schwein., Consp. Fung. Lusat.: 274, 1805.
BASIDIOCARPS annual, hard-coriaceous, resupinate, totally prostrate with
free margins 3-4 mm wide, in appearance as several small, hardly separable,
round-ovoid patches scattered on wood substratum, confluent and expanding
up to 5 cm in length. ABHYMENIAL SURFACE strigose-tomentose, easily
zonate, white-cream, with thin inflexed margin. HYMENIAL SURFACE even to
tuberculate, grayish-brown with violaceous tinge mostly in the central part of
Fic. 14. Microscopic structures of Stereum sanguinolentum:
a — hyphae; b - hymenial fragment showing slender thick-walled pseudocystidia
among acanthohyphidia, basidia, and basidiospores. Scale bar = 10 um.
Stereum in Israel ... 123
the patches, color becoming gradually lighter towards the undulating margin.
Hymenophore of dried specimens bleeding when bruised. ConTExT thin,
light-colored, 0.5-1 mm thick, in cross-section a reddish-brown thin deposit is
seen (barely observable without magnifier).
HyPHAL SYSTEM pseudodimitic. Generative hyphae simple-septate,
hyaline, thin- to thick-walled, 3-4 um diam, branched; pseudoskeletal hyphae
thick-walled, rarely septate, 3-6 um diam. Conducting hyphae of the same
dimensions, but without septa and with oil-rich brown contents, occur in
subhymenial and medullar tissues. CysTrp1 not seen but cystidia-like hyphal
ends (pseudocystidia) present. PsEUDOCYSTIDIA cylindrical to subcylindrical,
some with an inflated middle part up to 13 um in diam., and narrow base; with
basal walls 1.5-2 um thick and brownish oily contents. ACANTHOHYPHIDIA
abundant, cylindrical, tapering towards apex, thin-walled, 20-30 x 3-4 um
in diam. Basrp1a cylindrical to narrowly clavate, 25-55 x 3-10 um, 4-spored,
without a basal clamp. Basip1osPorss ellipsoid to cylindrical, hyaline, smooth,
thin-walled, (6-)7-10(-10.5) x (2—-) 2.5-3(-3.5) um, amyloid.
The species was found inhabiting pine wood in North of Israel (Fie. 15).
GENERAL DISTRIBUTION AND HABITAT. Being associated with a white rot,
Stereum sanguinolentum is a very affective and fast colonizer of newly dead or
wounded conifer sapwood, probably the most important fungus involved in
“wound rot of spruce” (Schmidt 2006, Calderoni et al. 2003).
Many authors reported this species as parasitized by jelly | »
fungus Tremella encephala Pers. According to our knowledge, —
the general distribution of Stereum sanguinolentum includes:
Europe (Austria, Czech Republic, Denmark, Finland, France, *
Georgia, Germany, Iceland, the Netherlands, Norway, Portugal,
Romania, Russia, Slovenia, Spain, Sweden, UK, Ukraine);
MippLE East (Israel); East Asta (Japan, Mongolia, South
Korea); NORTH AMERICA (Canada, USA); OTHER (Faeroe
Islands).
Fic. 15. Distribution of Stereum sanguinolentum in Israel.
Note. A notable difference distinguishing S. sanguinolentum from other
similar species in the field is its coniferous substrate. Due to its red staining
reaction when bruised, S. sanguinolentum might be confused with S. rugosum,
S. gausapatum, and representatives of the S. hirsutum-complex. Diagnostic
differences are: substratum, spore size, and presence of colored conducting
hyphae. Jiilich & Stalpers (1980) reports that although acanthohyphidia are
mentioned in descriptions of American specimens, they are never reported for
European material. The studied Israeli material is characterized by numerous
acanthohyphidia.
124 ... Tura & al.
SPECIMEN EXAMINED: ISRAEL. GH, near Mt. Baron, Pinus forest, on pine fallen wood,
26.11.2006, leg. D. Tura, det. D. Tura, I. Zmitrovich et V. Malysheva (HAI 0143).
Stereum subpileatum Berk. & M.A. Curtis, Hook. J. Bot. Kew Gard. Misc. 1: 238,
1849. Fic. 16
= Xylobolus subpileatus (Berk. & M.A. Curtis) Boidin, Rev. Mycol. 23: 341, 1958.
“Sporophore effused-reflexed, attached by the umbos, small, 1-1.5 cm long and
up to 1.5 mm thick; pileus tomentose, grayish-orange to light brown or brown,
somewhat zonate; margin brownish; hymenial surface light orange, smooth or
somewhat pubescent. Context pale-orange. Hyphal system trimitic; skeletal
hyphae thick-walled, 4-7 um wide. Cystidia cylindric, thick-walled, encrusted,
5-7 um wide. Basidia 20-40 x 3-5 um, with 4 sterigmata; spores cylindric,
4-4.5 x 2-2.5 um, hyaline, smooth” (Binyamini 1982).
The hyphal system of this species can be characterized as pseudodimitic to
subdimitic (pseudoskeletal hyphae in older parts lose their plasmatic contents).
Some sclerified elements are actually branched similar to “trimitic” polypore
elements, a condition also observed in other Stereum species.
GENERAL DISTRIBUTION AND HABITAT. Associated with white pocket rot,
Stereum subpileatum occurs on living hardwood substrata, mainly oak. Its
general distribution includes Europe (Austria, Macedonia, Russia, Spain);
Mipp1E East (Israel); East Asta (China, Japan, Nepal); NorTH AMERICA
(Canada, Cuba, Mexico, USA); SouTrH AMERICA (Colombia, Costa-Rica,
Venezuela).
Note. According to Binyamini (1982), this species was found
in Israel only once, in Samaria (Fic. 16), and differs from | .
other Israeli Stereum species by widely effused, hard, perennial || *
fruitbodies with cylindrical encrusted cystidia, and by the hard
cover of small pilei (see also Chamuris 1988, Julich & Stalpers -
1980). Microscopically, Stereum subpileatum is highly similar
to S. frustulatum (Pers.) Fr. (both considered conspecific by K~« Ks
Welden, 1971). In many handbooks the two species have been | \"
treated as separate species belonging to the genus Xylobolus.
However, we prefer another solution (see Introduction).
Fic. 16. Distribution of Stereum subpileatum in Israel.
Stereum in Israel... 125
Key to Israeli species of Stereum
PoieAcanthOny puldia AUCCIt Wate eam mn EN sree tore kM os alg os eck gle ae p!
Pee ACANUNOny phidia DICSe ll tk aera rere eR eee las acs crak ance ee 3
2a. Basidioles cylindrical. Pseudocystidia basal walls equal/less than 1.5 um thick.
Abhymenial surface condensed, subtomentose to nude. Hymenium bleeding
when bruised, with crysescent conducting hyphae) ............. S. gausapatum
2b. Basidioles acuminate-cylindrical. Pseudocystidia thick-walled, basal walls usually
more than 2 «um thick. Abhymenial surface hirsute-tomentose. Hymenium when
injured, in basidiocarps with active growth, sometimes staining yellow-orange-
PCC DUteWITNOUL CONGUCHINO LY Didceerta twee st aie Oeste S. hirsutum
3a. Basidiocarps annual, tough-coriaceous, not deeply cracking. Hymenium bleeding
when injured, with abundant conducting hyphae. Pseudocystidia basal walls
1.5-2 um wide. Basidiospores (6-)7-10(-10.5) x 2.5-3(-3.5) um
5 D0, St A RSPR PPE “OR ok SRR aa Nar ORT S. sanguinolentum
3b. Basidiocarps perennial, hard-corneous, with deeply cracked multistratose
hymenium. Hymenium not bleeding when injured, without conducting hyphae.
Basidiospores:449 902-7, (Ul eae pene oe yas te Eee S. subpileatum
Acknowledgments
The authors are very grateful to Dr V.E Malysheva (Russia) for help during field
researches and granting a photo, Prof. R.H. Petersen (USA), Dr W.A. Spirin (Russia),
Dr. S.R. Pennycook (New Zealand), and Dr. L.L. Norvell (USA) for critical review of the
manuscript and valuable remarks. The research was partly supported by RFFI grant (NN
06-04-49043, 06-04-49524, 07-04-01408).
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126 ... Tura & al.
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MYCOTAXON
Volume 106, pp. 127-132 October-December 2008
A note on Stiptophyllum erubescens
MARISA DE CAMPOS-SANTANA”?! & CLARICE LOGUERCIO-LEITE!
*marisacampossantana@yahoo.com.br
‘Laboratorio de Micologia - Depto. Botanica - PPGBV/CCB
Universidade Federal de Santa Catarina
Campus Universitario, 88090-040, Florianopolis, Santa Catarina, Brazil
Abstract — Fresh basidiomata of Stiptophyllum erubescens were studied and its macro
and microstructures, mainly basidia and basidiospores, were elucidated.
Key words — Agaricomycetes, Polyporaceae, taxonomy
Introduction
Stiptophyllum Ryvarden (Polyporaceae, Basidiomycota) was described in 1973
with Daedalea erubescens as type species. With its stipitate basidiomata, it
was proposed as a satellite genus to Gloeophyllum P. Karst., which has sessile
basidiomata. Stiptophyllum is macroscopically distinguished by stipitate
basidiomata, a tomentose pileus and stipe, and a lamellate hymenial surface.
Microscopically it is characterized by a trimitic hyphal system and cylindrical,
hyaline, and non-amyloid basidiospores (Ryvarden 1973). The genus is
monotypic, causes a brown rot, and is endemic to tropical South America.
Basidiospores and basidia are notoriously difficult to observe in old and/or
dried basidiomata since they apparently collapse very rapidly. The present study,
carried out with fresh material, made it possible to elucidate these structures
more accurate than earlier ones (Fidaldo 1959, Fidalgo 1968, Ryvarden 1973,
Singer 1975, Wright & Deschamps 1977).
Materials and methods
Basidiomata of S. erubescens were collected during a recent field survey in the
municipality of Mondai (27°0616’S and 53°24”07’W) in Linha Uruguai and
Linha Sanga Forte, in the extreme west of the State of Santa Catarina in southern
Brazil. The area was originally covered by Deciduous Seasonal Forest, and the
samples for this study were collected in remnants of this original vegetational
128 ... Campos-Santana & Loguercio-Leite
type (Ruschel et al. 2003). Macro- and microscopical data of the specimens were
obtained following the traditional methodology (Singer 1975, Ryvarden 1991),
and the colors were determined according to Munsell (1975). The examined
specimens are preserved in Herbarium FLOR (Holmgren & Holmgren 1998).
Taxonomy
Stiptophyllum erubescens (Berk.) Ryvarden, Norweg. J. Bot. 20: 4.1973. FIGS 1-2
=Daedalea erubescens Berk., Ann. Nat. Hist. 4: 292. 1840.
=Lenzites erubescens (Berk.) Sacc., Syll. Fung. (Abellini) 5: 645. 1887.
=Cellularia erubescens (Berk.) Kuntze, Revis. gen. pl. (Leipzig) 3(2): 452. 1898.
=Xerotus erubescens (Berk.) Singer, Los Angeles County Mus. Contr. Sci. 24: 6. 1958.
=Xerotinus erubescens (Berk.) Singer, Sydowia 15: 83. 1962 ['1961’].
=Gloeophyllum erubescens (Berk.) Popoff, Mycotaxon 87: 105. 2003.
=Lenzites guilleminiana Lév., Ann. Sci. Nat., Bot., sér. 3, 5: 122. 1846.
=Lentinus guilleminianus (Lév.) Mont., Ann. Sci. Nat., Bot., sér. 3, 5: 364. 1846.
=Pocillaria guilleminiana (Lév.) Kuntze, Revis. gen. pl. (Leipzig) 2: 866. 1891.
=Lentinus schomburgkianus Henn., Hedwigia 36: 205. 1897.
=Lentinus frondosus Henn., Hedwigia 36: 206. 1897.
=Lentinus fuscoferrugineus Speg., An. Mus. nac. Hist. nat. B. Aires 19: 226. 1909.
BASIDIOMATA annual, laterally or eccentrically stipitate, solitary or in groups,
connected at the base of the stipes or sometimes in groups with entirely free
stipes and laterally fused pilei. PrLEUs rounded to almost infundibuliform or
flabelliform, with a large variation in size, cartilaginous, flexible and somewhat
humid when fresh, firm and light in weight when dry, up to about 15 x 12.5 x
1.5 cm; pileus surface trichodermoid, without cuticle, coriaceous when young
becoming rigid and woody in time, finely subtomentose to glabrous, rough,
concentrically zonate and sulcate, striate near to the margin, yellow (8/6-8/8,
7/6-7/8 1OYR) to brownish yellow (6/6-6/8 10YR) and dark yellowish brown
(3/6 1OYR) when dry; margin irregular, acute, rigid, undulate or lobate, white
(8/1 5Y) when fresh becoming pale brown (6/3 10YR) to dark brown (3/3 10YR)
when dried. Stripe subwoody to woody, initially velutinous then scabrous and
longitudinally sulcate, 1-6 x 1.5-4.5 cm, very pale brown (7/4 10YR) when
fresh to dark yellowish brown when dried (4/4-3/4 10 YR), pseudo-sclerotium
present at the base of the stipe, always wider than the rest of the stipe.
HyMENOPHORE lamellate, subdaedaloid or irpiciform, lighter than the surface
of the pileus, very pale brown (7/4 10YR) when fresh to light yellowish brown
to brownish yellow (6/4-6/8 10YR) when dry, lamellae coriaceous to rigid,
deeply decurrent, 6-12 per cm at the margin, edges initially entire then crenate,
irregular, up to 5 cm in length and 0.9 cm deep, very pale brown when fresh
(7/4 10YR) to brownish yellow (6/6 10YR), when dry become dark yellowish
brown (4/6 10YR). CONTEXT corky, conspicuous, homogeneous, contrasting
with the pileus surface, soft, almost tough when dry, 1-4 mm thick, brownish
yellow (6/8 1OYR) to dark yellowish brown (3/4 10YR).
Styptophyllum erubescens ... 129
FiGuRE | - General aspect of basidiomata: FLOR 32202, 32203, 32204 and 32206.
Scale=5cm:
HYPHAL SYSTEM - trimitic; in the trama generative hyphae hyaline, clamped,
thin-walled to slightly thick-walled, 2-3(-4) um in diam.; skeletal hyphae
yellowish to brown, unbranched, non-septate, thick-walled to solid, tortuous,
walls with many constrictions, sometimes with moniliform aspect, 3-7 um in
diam.; binding hyphae subhyaline to yellowish brown, very branched, 2-4 um
diam., thick-walled to subsolid. Hyphal system of the context similar to the
trama, in stipe hyphae slightly dextrinoid, simple-septate. Basip14 hyaline to
slightly stained, clavate and tetrasporic, thin-walled and smooth, constricted or
sometimes collapsed at the base, 35-40 x 5-6 um; basidioles smaller than the
basidia, 20-33 x 2-4 um; Basip1osPorREs cylindrical-ellipsoid, hyaline to slightly
yellowish, thin-walled, 9-12 x 3—4(-5) um, inamyloid and nondextrinoid.
SUBSTRATE — dicotyledonous dead wood.
DISTRIBUTION — Neotropical; Bolivia, Colombia, Peru, Venezuela, Guiana,
Argentina, Paraguay and Brazil: Acre, Rio Grande do Norte, Rio de Janeiro,
Mato Grosso, Rio Grande do Sul, Amazonas, Para, Goidas, Bahia, Paraiba,
Parana and Santa Catarina (Fidalgo 1968, Ryvarden 1973, Ryvarden 1991,
Singer 1975, Wright & Deschamps 1977, Bononi 1992, Ryvarden & Iturriaga
2001, Popoff 2003, Gibertoni et al. 2004, Drechsler-Santos 2005, Campos-
Santana & Loguercio-Leite 2008).
130 ... Campos-Santana & Loguercio-Leite
FiGuRE 2 — A. Hymenium. B. Generative hyphae from stipe. C. Basidiospores.
D. Skeletal hyphae from trama. E. Binding hyphae from trama. F. Generative hyphae.
Scale =10 um.
VOUCHER MATERIAL — BRASIL, SANTA CATARINA: Mondai 15/IV/06 Linha Sanga
Forte, Campos-Santana & Santana 71 (FLOR 32202), Linha Sanga Forte, Santana &
Zanella 91 (FLOR 32203); 23/V/07 Linha Sanga Forte, Campos-Santana & Santana 236
(FLOR 32304), 256 (FLOR 32205) & 297 (FLOR 32206).
ADDITIONAL MATERIAL — BRAZIL, Santa Catarina: Itapiranga 15/IV/1985 Linha
Becker, Scholz (FLOR 10108).
REMARKS. Specimens of Stiptophyllum erubescens are flexible and moist
when fresh, firm and lighter in weight when dry. The subwoody to woody
stipe is initially velutinous then scabrous and longitudinally sulcate and has’
a pseudosclerotium at the base. As noted by Wright & Deschamps (1977),
specimens stain dark brown when touched with KOH. Skeletal hyphae in the
stipe appear slightly dextrinoid and generative hyphae are simple-septate, both
features cited for this species here for the first time. In previous cultural studies,
Wright & Deschamps (1977) described brownish simple-septate hyphae that
Styptophyllum erubescens ... 131
contrast with observations by Ryvarden (1973), “...binding hyphae hyaline,
thick-walled to solid, some ending in the hymenium as solid cystidia..., which
were not seen in our specimens. In some specimens (FLOR 32204; 32206)
the basidiospores - generally considered important for classification of the
Agaricomycetes (Ryvarden 2004) - were ellipsoid to cylindrical [9-12 x 3-4
(—5) um], hyaline to slightly brown, thin-walled, inamyloid and nondextrinoid.
Wright & Deschamps (1977), however, reported smaller basidiospores [6.6-8.2
(-9.6) x (1.8-)2.6-3.6 um]. Neither Ryvarden (1973) nor Drechsler-Santos
2005) cite basidiospore size data, although Ryvarden (2004) reported that
specimens collected during periods without active growing have few or no
basidiospores. We also did not find basidiospores in some specimens (FLOR
32202, 32203 and 32205). Basidia sizes [35-40 x 5-6 um] in our specimens are
similar [20-52 x 4.1-6.5 um] to those cited by Wright & Deschamps (1977) but
are larger than those reported by Fidalgo (1968) [16.5-24 x (4-)4.5-7.5 um].
The taxonomic position of S. erubescens is still subject to debate. Singer,
who accepted it in Xerotus Fries, initially stressed the presence of stipe as the
main difference separating it from Gloeophyllum (Singer 1975) but later added
the hyphal system as also diagnostic (Singer 1986). Fidalgo (1968) and Wright
& Deschamps (1977) both considered S. erubescens to have a dimitic hyphal
system. As previously mentioned, Ryvarden (1973) based Stipitophyllum on the
presence of a stipe and trimitic hyphal system. Recently, Popoft (2003) proposed
a new combination — Gloeophyllum erubescens, placing the main emphasis on
the brown rot decay. Most recently, Hibbett et al. (2007) proposed the order
Gloeophyllales Thorn to accommodate all species in the Gloeophyllum clade
(Binder et al. 2005), which includes species producing brown rot (Gloeophyllum,
Neolentinus, Veluticeps) or stringy white rot (Boreostereum, Donkioporia). No
molecular studies have yet been performed on S. erubescens (Kim & Jung 2000,
Binder et al. 2005) but future sequence analyses may help clarify the taxonomic
situation of this species and determine whether it belongs to an independent
genus or is part of Gloeophyllum.
Acknowledgments
We are grateful to Leif Ryvarden (Norway), Aristételes Gdes-Neto (Brazil), and Shaun
Pennycook (New Zealand) for their help in the correction and improvement of the
present paper. This work is part of M.Sc. thesis in Biologia Vegetal (Universidade Federal
de Santa Catarina, Brazil) of the first author.
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Munsell L. 1975. Munsell soil color charts. United States Department of Agriculture Hand. 18. Soil
Survey Manual.
Popoff OF. 2003. Notes on Daedalea erubescens, Hexagonia decipiens and the Phaeotrametaceae.
Mycotaxon. 87: 103-110.
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da Floresta Estacional Decidual do Alto-Uruguai, SC. Ciéncia Florestal.13: 153-166.
Ryvarden L. 1973. New Genera in the Polyporaceae. Norwegian Journal of Botany 20: 1-5.
Ryvarden L. 1991. Genera of Polypores - Nomenclature and taxonomy. Synopsis Fungorum 5:1-
363.
Ryvarden L, Iturriaga T. 2001. Studies in Neotropical Polypores 9. A Critical Checklist of Poroid
Fungi from Venezuela. Mycotaxon. 78: 393-405.
Ryvarden L. 2004. Neotropical polypores Part 1. Synopsis Fungorum. 19: 1-229.
Singer R. 1975. The Agaricales in Modern Taxonomy. 3rd ed. J. Cramer: Vaduz (Liechtenstein).
Singer R. 1986. The Agaricales in Modern Taxonomy. 3th ed. Koeltz: Koenigstein.
Wright JE, Deschamps JR. 1977. Basidiomicetos xilofilos de la Region Mesopotamica III. Los
Géneros Bjerkandera, Gloeophyllum, Gloeoporus, Hirschioporus, | Hydnopolyporus,
Phaeocoriolellus, Pycnoporus y Xerotinus. Buenos Aires, Revista de Investigaciones
Agropecuarias INTA 5 13: 27-70.
MYCOTAXON
Volume 106, pp. 133-178 October-December 2008
Taxonomic studies on Ustilaginomycetes - 28
KALMAN VANKY
vanky.k@cityinfonetz.de
Herbarium Ustilaginales Vanky (HUV), Gabriel-Biel-Str. 5
D-72076 Tiibingen, Germany
Abstract—The problem of generic delimitation of Tranzscheliella and Tranzscheliella-
like smut fungi is discussed. New species described are: Sporisorium mandlaicum,
S. shivasiorum, Thecaphora arnicae, Tranzscheliella poae, Ustilago milii-vernalis,
U. sclerachnes. New name proposed: Sporisorium ciccaronei. New combinations
proposed: Bauerago tinantiae, Entyloma arnaudianum, Gjaerumia eremuri, G. muscari,
Sporisorium stiparum, Thecaphora solidaginis, Tolyposporium isolepidis, Tranzscheliella
distichlidis, Tr. iranica, Tr. stipae-barbatae, Urocystis bulbinellae, Ur. simplex. Taxa placedin
synonymy are: Entyloma crepidis-tectorum (= E. zacintha), E. lavrovianum (= E. hieracii),
Urocystis radicicola (= Ur. rechingeri), Ustilago duriusculae (= U. striiformis) and
Ustilago moehringiae (= Microbotryum duriaeanum). Excluded from the smut fungi
is: Melanotaenium spermacoces. Lectotypes are designated for Entyloma lobeliae and
Sorosporium solidaginis. Keys are given to the species of Gjaerumia, of Melanotaenium,
and of Tranzscheliella, to the smut fungi of Chionachne and related genera, of Elionurus,
of Eulalia and Eulaliopsis, and of Sehima, to the Urocystis species of Anemone and
Anemonella, and to the genera of smut fungi.
Key word—taxonomy
Tranzscheliella and Tranzscheliella-like smut fungi
Lavrov (1936:29) described a new genus and species, Tranzscheliella otophora
Lavrov (= T: williamsii), on the basis of two, small cells attached at opposite
sides of the spores of a smut fungus collected on the stems of Stipa pennata
from Turkmeniya. Only half a century later, with help of a scanning electron
microscope (SEM), it was demonstrated (Vanky 1985:256, Fig. 240; 1987:121;
1994:446, Figs. 466a, b, c) that these two small cells were actually the oppositely
situated opercula or lids, which are circularly broken parts of the thick exospore,
attached to the spore on a relatively short part of their circumference. The
opercula are closed when the spores are dried, and opened when the spores
are rehydrated. Presence of opercula alone did not seem to be a sufficiently
strong morphological character for recognising a separate genus. Together
with the morphology of sori, the spores as well as spore germination, the genus
134 ...Vanky
Tranzscheliella was recognised as a well circumscribed group of smut fungi,
all on members of Poaceae (Vanky 2002:162). Because of the morphological
similarities between Ustilago hypodytes and Tranzscheliella williamsii (except
for the opercula), U. hypodytes was transferred into this genus as 1. T: hypodytes
(Schltdl.) Vanky & McKenzie (Vanky & McKenzie 2002:156; type on Elymus
arenarius L., Germany). Several smut fungi, with similar morphology, i.e.
possessing external, naked, powdery sori on the surface of the stems, rarely
also on floral axes of Poaceae, and solitary, brown, small-sized spores, were
also transferred into the genus Tranzscheliella (Vanky 2003:2-4, 2004:109),
or described as new species. These are: 2. T. amplexa (Syd.) Vanky (Vanky
2004:109; type on Diplachne fusca (L.) P. Beauv. ex Roem. & Schult., Egypt),
3. T. austrostipae R.G. Shivas & Vanky (Shivas & Vanky 2007:9; type on
Austrostipa verticillata (Nees ex Spreng.) S.W.L. Jacobs & J. Everett, Australia),
4. T. comburens (F. Ludw.) Vanky & McKenzie (Vanky & McKenzie 2002:155;
type on Stipa sp., = misnamed Danthonia sp., Australia), 5. T. halophila (Speg.)
Vanky (Vanky 2003:2; type on Distichlis scoparia (Kunth) Arechav., Argentina),
6. T. halophiloides (G.W. Fisch.) Vanky (Vanky 2003:2; type on Distichlis stricta -
(Torr.) Rydb., USA), 7. T. jacksonii (Zundel & Dunlap) Vanky (Vanky 2003:3;
type on Stipa lettermanii Vasey, USA), 8. T. laevispora Vanky (Vanky 2004:82;
type on Sporobolus agrostoides Chiov., Kenya), 9. T: macrochloae (Pat.) Vanky
(Vanky 2003:3; type on Macrochloa tenacissima (L.) Kunth = Stipa tenacissima
L., Tunisia), 10. T. minima (Arthur) Vanky (Vanky 2003:3; type on Oryzopsis
cuspidata Benth. ex Vasey, USA), 11. T. serena (Syd.) Vanky (Vanky 2004:109;
type on Diplachne fusca, Australia), 12. T. sparti (Massenot) Vanky (Vanky
2003:4; type on Lygeum spartum L., Tunisia), 13. T. ventanensis (Hirschh.)
Vanky (Vanky 2003:4; type on Stipa sp., Argentina), and 14. T. williamsii
(Griffiths) Dingley & Versluys (Dingley & Versluys 1977:477; type on Stipa
richardsonii Link, USA).
In 2002, mycologists from University of Tubingen and I started a detailed
study of the genus Tranzscheliella, including molecular analyses. An
important taxonomic conclusion from the interrupted study was that not all
morphologically Tranzscheliella-like smut fungi belong to this genus. This
became evident by the analysis of T: comburens, which actually belongs to the
genus Ustilago, as originally described (M. Stoll & D. Begerow, pers. comm.).
Further Tranzscheliella-like smuts, U. triodiae Vanky (comp. Stoll et al. 2005)
and U. lituana R.G. Shivas et al., are also true Ustilago species (comp. Shivas et
al. 2006:365).
Recently, Begerow et al. (2007:914) demonstrated by molecular phylogenetic
analyses that “Macalpinomyces and Tranzscheliella, which both occur on
Poaceae, are not members of the clade representing Ustilaginaceae.’ The isolated
position of Tranzscheliella outside the large Ustilago-Sporisorium complex may |
Ustilaginomycetes — 28 ... 135
justify its recognition as a good genus. In the absence of reliable morphological
characters for the genus Tranzscheliella, molecular phylogenetic analyses may
be necessary to resolve species delimitation. It appears that Tranzscheliella
hypodytes, T: minima, T. sparti and T: williamsii are true species (M. Stoll &
D. Begerow, pers. comm.). In the absence of molecular phylogenetic studies,
placement of some smuts into Tranzscheliella or Ustilago is uncertain and
can be considered only provisional. Similarly, uncertain are several of the
synonyms and the host plant genera of T’ hypodytes s.lat., enumerated in Vanky
& McKenzie (2002:156-158).
Molecular phylogenetic analysis of Tranzscheliella “hypodytes” on Poa cita from
New Zealand showed an isolated position from all other analysed Tranzscheliella
species (M. Stoll & D. Begerow, pers. comm.). This means that the smut on
Poa cita represents a separate species. If all T’ hypodytes-like smuts on various
Poa species were conspecific, the name Ustilago custanaica Lavrov (type on
Poa pratensis, Siberia, deposited where?) could be applied to them. Two of
the collections on Poa species, namely Poa poiformis from Australia (HUV
17630) and Poa nevadensis from the USA (HUV 10169), have spores that are
morphologically identical with those of T: hypodytes (spores 4—6.5 um long, with
polar caps), whereas those of the analysed Poa cita have smaller spores (3.5-5.5
tum long, only with polar thickenings). The spores in the original description of
Ustilago custanaica are 3.5-7 «1m long, and it has to be considered a synonym of
T: hypodytes. The smut on Poa cita represents a new species.
Tranzscheliella poae Vanky, sp. nov.
MycoBank: 511638
Typus in matrice Poa cita, New Zealand, South Island, Central Otago, 20 km S oppid.
Alexandra, Old Man Range, Symes Road, 45°20°17” S, 169°17°04” E, alt. 750 m.s.m.,
6.11. 1998, leg. E.H.C. McKenzie, C. & K. Vanky. Holotypus in Herbario Ustil. Vanky, HUV
18620, isotypi in PDD 69200, et in Vanky, Ust. exs. no. 1049 (ut Ustilago hypodytes).
Paratypus in matrice Poa cita, New Zealand, South Island, Mid-Canterbury, Christchurch,
Port Hills, Victoria Park, 13.XII.1981, leg. G.C. Kirby, PDD 41990, isoparatypus in HUV
16473.
Tranzscheliella poae differt a specie T. hypodytes (Schitdl.) Vanky et McKenzie, Smut
Fungi of New Zealand:156, 2002. (Caeoma hypodytes Schltdl., Fl. Berol., Pars 2.
Cryptogamia:129, 1824) sporis minoribus (3-4,5 x 3,5-5,5 um), capsulis polaribus
minoribus, incrassationibus tantum laterum deplanatorum, et imprimis characteribus
biologicis molecularibus distinctis. Sporae speciei T. hypodytes 3,5-5,5 x 4-6(-7) um,
plerumque pileo polari hyalino minuto instructae.
Sori (Fig. 1) surrounding the distal internodes of sterile shoots, more or less
hidden by leaf sheaths, spore mass naked, dark brown, powdery. Sporgs (Figs.
4, 5) globose, subglobose, slightly flattened, 3-4.5 x 3.5-5.5 um, olivaceous-
brown; wall even, c. 0.5 um thick, in LM apparently smooth to very finely
punctate, often with two flat, hyaline, finely verruculose thickenings, i.e. weakly
136 ...Vanky
natn tein iuccinnnonicnsisin z
cin a tessa Sonata oRe rs resecpat <
irentsetetre
(type). Habit, and to the left a healthy
Fig. 1. Sori of Tranzscheliella poae on the stem of Poa cita
Bar= lems
inflorescence.
Ustilaginomycetes — 28 ... 137
developed polar caps on the flattened sides, in SEM finely, rather densely
provided with warts of variable sizes due to fusion.
On Poaceae: Poa cita Edgar (P. caespitosa Spreng.); New Zealand.
Tranzscheliella poae differs from T. hypodytes in having smaller spores, weakly
developed polar caps, and especially by different molecular biological characters
in T. poae (unpublished data). In the literature, several Poa species are given as
host plants of “Ustilago” hypodytes, including P. nevadensis Vasey, P. poiformis
(Labill.) Druce, P. pratensis L., PR ampla Merr., PB. canbyi (Scribn.) Howell,
P juncifolia Scribn., P. palustris L., and P. scabrella (Thurb.) Benth. (P. buckleyana
Nash), mostly from N America but also from E Asia. Some of these hosts could
harbour T° poae.
Because of the close resemblance of the character complex of the sori and
spores to the type species Tranzscheliella hypodytes, and also using host plant
criteria, three further smut fungi are considered best placed into the genus
Tranzscheliella. These are:
Tranzscheliella distichlidis (McAlpine) Vanky, comb. nov.
MycoBAnk: 511639
BASIONYM: Cintractia distichlidis McAlpine, The smuts of Australia:169, 1910, as
‘distichlydis. — Ustilago distichlidis (McAlpine) Cif, Ann. Mycol. 26:32, 1928, as
‘distichlydis. — Lectotype on Distichlis maritima (= D. distichophyila), Australia,
(design. by Vanky 2001:321) Victoria, Port Phillip, X.1891, C. French jr, MEL 1055118};
isolectotypes HUV 19156!, VPRI 2923. Paratype on D. maritima, Australia, Victoria,
Sandringham, 5.[X.1905, D. McAlpine, VPRI 2924.
SORI surrounding the distal internodes of sterile shoots, more or less hidden
by the leaf sheaths, at first covered by a thin, greyish peridium that gradually
flakes away, exposing the dark brown, powdery mass of spores. SPORES globose,
subglobose, ellipsoidal, 4.5-6 x 5-6.5 um, reddish-brown; wall even, c. 0.5 um
thick, polar caps absent, in LM smooth, in SEM finely, sparsely to moderately
densely verruculose.
On Poaceae: Distichlis distichophylla (Labill.) Fassett (D. maritima Raf.; D. spicata auct.
non (L.) Greene); Australia.
Tranzscheliella iranica (Syd.) Vanky, comb. nov.
MycoBank: 511640
BasionyM: Ustilago iranica Syd., Ann. Mycol. 37:199, 1939. — Type on Stipa tenerrima,
North Iran, Elburs Mountains, Mt. Kuh-e-Nemar, near Panshand, 3. VI.1937, D.E. Gauba
1233. Holotype B, isotype HUV 14817!
Sori destroying the whole inflorescence, partly enclosed by the uppermost
leaf sheaths, up to 7 cm long, with black or blackish-brown, powdery spore
mass, peridium absent. Spores subglobose, broadly ellipsoidal or rounded
subpolyhedrally slightly irregular, 4-7 x 5-8 um, yellowish- to olivaceous-
138 ...Vanky
brown; wall 0.5-1.5 um thick, from apparently smooth to finely, densely
verrucose, spore profile smooth to finely wavy, no polar caps, in SEM with
rounded, low warts fusing into irregular groups or short, irregular rows.
On Poaceae: Stipa tenerrima Bornm. & Gauba; Asia (Iran). Known only from the type
locality.
Tranzscheliella iranica is close to the South American T: ventanensis, in which
the spores are pronouncedly verrucose and the sori surround the uppermost
internodes and aborted inflorescence. The report of T: iranica from China (Guo
2000:73) may in fact represent a Sporisorium sp.(?) or result of insect infestation
(L. Guo, pers. comm.).
Tranzscheliella stipae-barbatae (Maire) Vanky, comb. nov.
MycoBank: 511641
BastonyM: Ustilago stipae-barbatae Maire, Bull. Soc. Hist. Nat. Afrique N. 8:139,
1917. — Cintractia stipae-barbatae (Maire) Maire, in Maire & Werner, Mém. Soc. Sci.
Nat. Maroc. 45:44, 1937. — Lectotype on Stipa (design. by Vanky 1985:256) barbata,
Morocco, Taourirt, 4.VI.1916, L. Ducellier, Herb. Maire 4226, MPU! Paratype on Stipa
gigantea, Algeria, Miliana, near the top of Zaccar Rharbi, 12.VII.1917, R. Maire, Herb.
Maire 4810, MPU!
Sort in the whole inflorescence, enclosed by the uppermost leaf sheath, spore
mass naked, dark brown, powdery. Sporss globose, subglobose, ellipsoidal to
slightly irregular, slightly flattened (4—5.5 um wide), in plane view 5-6.5 x 5.5-7
(-8) um, dark brown, lacking polar caps; wall 0.8-1 um thick, thicker on the
flattened sides where, after boiling, indistinct opercula may appear, surface in
LM smooth, in SEM, at high magnification, finely, densely verruculose.
On Poaceae: Stipa barbata Desf., S. gigantea Link (Macrochloa arenaria (Brot.) Kunth),
S. pennata L.; N Africa, Asia.
Key to the species of Tranzscheliella
1. Spores in LM with 2, large, bipolar appendages, in SEM operculate. .T. williamsii
- SPOLeS TOL SOS Sav Attar ee eres tad he ar, gine oe ie Nee aE ts eee Zz
2. Spores in LM sparsely, evidently verrucose, in SEM with small tubercles
Ss SSRN an Os CUR eee PO ON cee ee Is aires Oe, eae een T: halophiloides
- SPOLes.NOUSO AA ache ae. eluate. Meech pea eee ee aay yok rare tee 8
3.4). Spores hemiglobose or cup-shaped jaw tue... setae ch ee eee eee 4
- Spores globose, subglobose, ellipsoidal or slightly irregular................... >
4. Spores 6-8(-9) tm long, in SEM very finely, densely verruculose. On Distichlis
Mt i Mea a A INR ee DR img gs Pe mT Labes 0 OMIA pe eh: T. halophila
- Spores 4.5-6.5 um long, in SEM smooth. On Sporobolus.......... T. laevispora
Jt pyoPOres. Oa. 5(— 10), Lin ONG areal whe anew eee ee T. jacksonii
- Spores Smaller a, = wwe ower lee Ae tes ete eke Mine ecient fey ee nen eee 6
Ustilaginomycetes — 28 ... 139
PES DOLCS@ a1 ro /1tir ly O10 Wan ere WEP ed Goud nine, Qs cia ss es u's siege a can aaeeme 7
- SSG INS SVC See eared cBiettiots S Scie 4 sa8 plc ceed ae toe ROR Rs re yr 8
y. Peridium present; spores in SEM sparsely verrucose. On Diplachne ....T. serena
- Peridium absent; spores in SEM densely verruculose. On Austrostipa
side be Te Mcisg cat ME AAR Res eat I aya le, Siem ia tan ghar an a aE rae T. austrostipae
8. Sorin tneunllorescence Wess e ene Se ees SOP EAE SGN, Wi Be naisd eae eo 9
~ SOMONE CULINS MEE TOR ine MEME eR o), mee ter ry ences bos soe OR ee 12
pam velleceveloped periciun present ume ayer ae: Ace ss T. macrochloae
- Peniditimiapsente haere: to. eee eo i, any eh a her eet Ain Sh 10
10. Spore wall 0.5-1.5 um thick, in SEM with rounded, low warts; fusing into short,
teres ular LOWS (O@eCTOUP SEO TSI ter mnt y veer An ce wie vt T. iranica
- Spore wall up to 1 um thick, in SEM finely, densely verruculose or indistinctly
11. Spores 6-8 um long, in SEM finely, densely verruculose. On Lygeum.... T. sparti
- Spores 5.5-7(-8) um long, in SEM indistinctly warty. On Stipa T. stipae-barbatae
PS es POLES 2 (6A Lin LOMO geme we geey niet ee Seti Be, Rs ot oe ve 13
~ Sporeslarcer pe reep ram (a etre. Pe Ue oo Series Sana etebactns wae 14
ommenvvell developed peridium presents a2). ao cee See eee ee T. minima
~ Peridiumabsenit a...) ewe eres Uy sor: Steichen, Aeeee SoZ Aer eee oe T. poae
14. Spores 5.5-8(-9) um long; warts in SEM confluent into irregular groups; sori on
thestemsiaidean theintlorescenccam ase) seni: sa ete eatineese sare T. ventanensis
- Spores smaller; warts not confluent into groups; sorion the stems ........... 15
15. A thin peridium present; spores 6-7(-7.5) um long; polar caps absent
eee et a hit icky eens Soc ae Ronee ss T. amplexa
~ Peridium absent; spores 4-6(—7) um long; small polar caps often present
er eee es ee as Seed T. hypodytes
New species
A new Sporisorium species on Sehima nervosum (Poaceae)
Sehima Forssk., a genus of five species in the Old World tropics, belongs to
the subfamily Panicoideae, tribe Andropogoneae, subtribe Ischaeminae. It has
been regarded as a segregate from Ischaemum (Clayton & Renvoize 1986:348).
On species of Sehima seven smut fungi are known: 1. Jamesdicksonia dactylidis
(Pass.) R. Bauer et al. (type on Dactylis glomerata L., Italy), 2. Sporisorium
nervosum Vanky et al. (type on Sehima nervosum (Rottler) Stapf, Australia),
3. S. queenslandicum Vanky et al. (type on Sehima nervosum, Australia),
4. S. sehimae (M.S. Patil) Vanky (type on Sehima nervosum, India),
5. S. sehimicola Vanky (type on Sehima ischaemoides Forssk., Zimbabwe),
6. S. sulcati (M.S. Patil) Vanky (type on Sehima sulcatum (Hack.) A. Camus,
India), and 7. Tilletia sehimae A.R. Patil et al. (type on Sehima nervosum,
India). A new species, collected recently in India is:
140 ...Vanky
son pstranetionenenoemscesanneoebeeteinen
ee osc peice nated
Fig. 2. Sori of Sporisorium mandlaicum in the inflorescence of Sehima nervosum (type). Habit.
Bar = 1 cm.
Fig. 3. Sori of Sporisorium ciccaronei on Elionurus muticus (type). Habit, and enlarged a healthy
spikelet, two infected spikelets and an infected inflorescence.
Bars = 1 cm for habit, and 2 mm for detail drawings.
Ustilaginomycetes — 28... 141
Sporisorium mandlaicum Vanky & N.D. Sharma, sp. nov.
MycoBAaNnk: 511642
Typus in matrice Sehima nervosum, India, Madhya Pradesh, Mandla, Kanha National
Park border, 1 km ab Turia gate, 22°07’ N, 80°26’ E, alt. cca. 700 m.s.m., 7.X.2006, leg.
N.D. Sharma. Holotypus in Herbario Ustil. Vanky, HUV 21400!, isotypi in BPI 877339 et
in HCIO.
Sporisorium mandlaicum a specie propinque Sporisorium queenslandicum Vanky et al.,
in Vanky e& Shivas, Fungal Diversity 7:159, 2001 (typus in matrice Sehima nervosum,
Australia) distinctum racemos plerumque partim destruens, sporis magis uniformibus,
majoribus, 8-10,5 x 8,5-11(-12) um, leniter sed magis conspicue echinulatis; sporae speciei
S. queenslandicum variables, 5,5-9 x 5,5-10 um, punctatae usque leniter echinulatae.
Sori (Fig. 2) destroying the raceme, often leaving intact some or many of the
distal spikelets, cylindrical or curved, 1-2 x 25-60 mm, at first covered by a
thick, brown peridium that flakes away and the dark brown, semiagglutinated
to powdery mass of loose spore balls, spores and sterile cells is scattered leaving
behind a long, simple columella, often with a few short lateral branches. SPORE
BALLS of many spores, ephemeral. Spores (Figs. 6, 7) globose, subglobose,
ovoid to broadly ellipsoidal, 8-10.5 x 8.5-11(-12) um, yellowish-brown; wall
even, c. 0.8 um thick, finely, densely echinulate, spore profile finely serrulate.
STERILE CELLS (Figs. 6, 7) in irregular groups, single cells variable in shape and
size, subglobose, ovoid, ellipsoidal or irregular with one or several flattened
sides, 5.5-17 x 6-21 um, subhyaline to pale yellow tinted; wall 0.5-0.8 um
thick, smooth, content homogeneous.
On Poaceae: Sehima nervosum (Rottler) Stapf, S Asia (India). Known only from the type
collection.
Sporisorium mandlaicum partially destroys the racemes, and has larger,
more uniform spores that are finely but more evidently echinulate than
S. queenslandicum. In S. queenslandicum the spores are variable, 5.5-9 x 5.5-10
[tm, punctate to finely echinulate.
Key to the smut fungi of Sehima species
1. Sori in the leaves as dark spots; spores embedded in the host tissue, not
DUIVeRUlen te gr mewnrn Naren mary ey er hee att, Jamesdicksonia dactylidis
~ S OrnnOminitheleavesuspOresspull VerUlent mere a. ae eee ren ee een tert eae 2
Pm Sorianithe ovaries, spores 30-3 71m in}diams.= eae Tilletia sehimae
- SOM GuilulicovaliessspOressm allt... «rt aes ere eee een eae ores, 3
3. Sori partially or totally destroying the inflorescence, several cm long.......... 7
* SOmmesiticted tone spikelets, less (nallel. > ClivONOtsme ne fete Cen mee reee 4
eee Spore pails ephemeral; sterile cells present... > 25 -- Sporisorium sehimicola
- Sporeballsratner persistent, sterile cellsalsenth =) emeren nese at een 5
142 ...Vanky
5. Soriin some sessile spikelets; columella 5-7; spore balls 50-160(—200) um
este rns en Ger) cane eer: era Aa ike BAL tc Oc Sporisorium sehimae
~ Sori in all sessile and pedicelled spikelets; columella 1; spore balls smaller...... 6
6. Spore balls 30-70(-100) um long; spore wall uneven, 1-2(-2.5) um, free surface
finely punctate se; 7. eee ee ee ae tet ee eee Sporisorium sulcati
- Spore balls 25-50(-55) um long; spore wall even, 0.5-0.8 um, free surface
COdTSely VELLUCUIOSC peeeiee 2 te mee eee wre ees Sporisorium nervosum
7(3). Sori destroying the whole inflorescence; spores 5.5-10 um long, punctate to finely
echinulatée vit dtatemnutin cn hee meee ia ce Sporisorium queenslandicum
- Sori often destroying partially the inflorescence; spores 8.5-11(-12) um long,
finely, denselveechinulate 2) | on veer eee Sporisorium mandlaicum
A new Sporisorium species on Elionurus muticus (Poaceae)
Elionurus Humb. & Bonpl. ex Willd., in the subfamily Panicoideae, tribe
Andropogoneae, subtribe Rottboelliinae has 15 species in tropical Africa and
America, and one species in Australia (Clayton & Renvoize 1986:362). On
Elionurus three smut fungi have been published: 1. Macalpinomyces elionuri-
tripsacoidis Vanky, type on E. tripsacoides Willd., Zambia, 2. Sporisorium
elionuri (Henn. & Pole-Evans) Vanky, type on E. muticus (Spreng.) Kuntze,
South Africa, and 3. S. elionuri-tristis Vanky, type on E. tristis Hack.,
Madagascar. Ciccarone (1951:224) described Sphacelotheca elionuri Ciccar. (as
‘elyonuri ) on Elionurus argenteus var. caespitosus (= E. muticus) from several
places around Asosa and Afodu, Ethiopia. Study of the syntypes revealed that
they represent a new species, which is described as:
Sporisorium ciccaronei Vanky, nom. nov.
MycoBank: 511643
REPLACED NAME: Sphacelotheca elionuri Ciccar., Mycopathol. Mycol. Appl. 5:224, 1951,
as ‘elyonuri, not Sporisorium elionuri (Henn. & Pole-Evans) Vanky. — Type on Elionurus
argenteus var. caespitosus (= E. muticus), Ethiopia, Benishangul-Gumuz Proy., “Asosa et
Afodu’, alt. c. 1600 m, HI.1939, coll. A. Ciccarone. Holotype BPI 195067!, isotype BPI
177497! Both nearly completely destroyed by insects.
Sori in spiculis inflorescentiae eiusdem nonnullis, ovoidei usque longe ellipsoidales,
0,5-1 x 2-3 mm, raro inflorescentiam totam occupantes, et tunc majores, 2-4 mm x
1-5 cm, primum peridio flavidobrunneo cooperti, quo rupto massam nigrobrunneam,
semiagglutinatam, granulosam usque pulveream glomerulorum sporarum laxarum, sporas
et cellulas steriles columellam crassam, decrescentem, conicam, 2-3 mm longam, sulcis
Figs. 4, 5. Spores of Tranzscheliella poae on Poa cita in LM and in SEM (type).
Figs. 6, 7. Spores and sterile cells of Sporisorium mandlaicum on Sehima nervosum in LM and in
SEM (type).
Figs. 8, 9. Spores and sterile cells of Sporisorium ciccaronei on Elionurus muticus in LM and in SEM
(type). Bars = 10 um.
Ustilaginomycetes — 28 ... 143
144 ...Vanky
longitudinalibus numerosis et nonnunquam cum ramis 1-2 apicularibus, spiniformibus
instructam cingentes. Sporae juveniles in grandibus, laxis, irregularibus glomerulis, in
statu maturo singulares, globosae, subglobosae, ovoideae, late ellipsoidales usque parum
irregulares, 6-8 x 6,5-9 um, rufobrunneae; pariete aequali, 0,8-1 um crasso, conspicue
levi usque subtiliter, dense punctato-verruculoso, imago obliqua sporae levis. Cellulae
steriles globosae, ellipsoidales usque irregulares, 8-16 x 11-18,5 um, hyalinae; pariete 2-3
um crasso, levi.
ETyMOLoGy: named in the honour of the excellent Italian phytopathologist and collector
of microfungi, Antonio S. Ciccarone (1909-1982), who also collected and described this
species.
Sor! (Fig. 3) in some spikelets of an inflorescence, destroying the inner floral
organs, often in groups, ovoid to long ellipsoidal, 0.5-1 x 2-3 mm, rarely
comprising the whole inflorescence, then larger, 2-4 mm x 1-5 cm, at first
covered by a yellowish-brown peridium that ruptures from its apex disclosing
the blackish-brown, semi-agglutinated, granular to powdery mass of loose
spore balls, spores and sterile cells surrounding a thick, narrowing, conical,
2-3 mm long columella with numerous longitudinal furrows and sometimes
with 1-2 spiniform, apical branches. Sporss (Figs. 8, 9) when young in large,
loose, irregular balls, when mature single, globose, subglobose, ovoid, broadly
ellipsoidal to slightly irregular, 6-8 x 6.5—9 um, reddish-brown; wall even, 0.8-1
um thick, apparently smooth to finely, densely punctate-verruculose, spore
profile smooth. STERILE CELLS (Figs. 8, 9) globose, ellipsoidal to irregular, 8-16
x 11-18.5 um, hyaline; wall 2-3 um thick, smooth.
On Poaceae: Elionurus muticus (Spreng.) Kuntze (E. argenteus var. caespitosus (A. Rich.)
Hack.); E Africa (Ethiopia).
Key to the smut fungi of Elionurus species (Poaceae)
1. — Sori in all spikelets of an inflorescence, long cylindrical; columella filiform;
Stetilescells abSen tates ae Rwrr oh Fim rn sa ered tc ae eevee Tat Sporisorium elionuri
- Sori in some spikelets or ovaries of an inflorescence, or comprising the whole
inflorescence; columella not filiform or absent, sterile cells present............ 2
2s Sori in some ovaries of an inflorescence, cylindrical; columella absent;
spores with short, conical warts .......... Macalpinomyces elionuri-tripsacoidis
- Sori in the spikelets or comprising the whole inflorescence; columella present;
Spores Otherwise Ornaln enter samen pen ater nee earner ee bere 2 ee 2
3; Sori in some spikelets of an inflorescence; columella conical, with longi-
tudinal furrows; spores 6.5-9 um long, apparently smooth to finely punctate-
verruculose; sterile cells 11-18.5 um long, wall 2-3 um thick
BF RMA WANS Sic ¢.% Egh PSP ER NR Ad hha A AOE Se Sporisorium ciccaronei
~ Sori in all spikelets of an inflorescence; columella short, simple; spores
10-13 um long, finely, moderately densely echinulate; sterile cells 7-13 um
lone, wWallice 0 Sain, Co lcke sr eh ter a ee eee Sporisorium elionuri-tristis
Ustilaginomycetes — 28... 145
A new Sporisorium species on Eulalia trispicata (Poaceae)
Eulalia Kunth, with c. 30 species in the Old World tropics, and Eulaliopsis
Honda, with two species from Afghanistan to China and the Philippines, belong
to the subfamily Panicoideae, tribe Andropogoneae, subtribe Saccharinae. Eulalia
is related to Saccharum. Eulaliopsis is a rather isolated genus, regarded as a
derivative of Eulalia (Clayton & Renvoize 1986:333). Eulalia species were often
treated under the generic name Pollinia Spreng., nom. rejic. (= Chrysopogon
Trin.). The smut fungi of Eulalia and Eulaliopsis were treated by me (Vanky
2000:183-186) and six species were recognised, all in the genus Sporisorium:
1. S. crypticum (Cooke & Massee) Vanky & M.S. Patil (type on Pollinia argentea
(Brongn.) Trin. = Eulalia trispicata, India), 2. S. eulaliae (L. Ling) Vanky (type
on Eulalia trispicata, Australia), 3. S. guangxiense L. Guo (type on Eulalia
pallens (Hack.) Kuntze, China), 4. S. polliniae (Magnus) Vanky (type on Pollinia
distachya (L.) Spreng., Israel), 5. S. pollinianum (Zundel) Vanky & R.G. Shivas
(type on Pollinia leptostachys Pilg., Papua New Guinea), and 6. S. indicum (Syd.
et al.) Vanky (type on Eulaliopsis binata (Retz.) C.E. Hubb., India). Recently, two
further species have been described: 7. S. trispicatae R.G. Shivas et al. (Vanky et
al. 2006:111; type on Eulalia trispicata, Thailand), and 8. S. rarum R.G. Shivas
et al. (Shivas et al. 2007:350; type on Eulalia aurea (Bory) Kunth, Australia). An
additional, new species, collected in Thailand, is described as:
Sporisorium shivasiorum Vanky, sp. nov.
MycoBAank: 511644
Typus in matrice Eulalia trispicata (det. H. Scholz, B), Thailand, Chiang Mai Prov., Mae
Taeng Distr., Mae Ngad Dam, 19°09’40.5” N, 99°02’24.1” E, alt. 397 m.s.m., 28.XII.2005,
leg. R.G., M.D.E., A.J., G.E Shivas et. al. Holotypus in Herbario Ustil. Vanky, HUV 21201],
isotypus in BRIP 51766.
Sporisorium shivasiorum a specie S. trispicatae R.G. Shivas et al., in Vanky et al.,
Mycol. Balcan. 3:111, 2006 (typus in matrice Eulalia trispicata, Thailand) ei arcte
cognato distinctum praecipue soris, glomerulis sporarum et morphologia sporarum. Sori
S. shivasiorum in omnibus ovariis inflorescentiae eiusdem, glomeruli sporarum
permanentes, sporae externae 9-12 um longae, mediocriter atro-flavidobrunneae, in
latero libero verrucis usque 0,4 um altis. Sori speciei S. trispicatae in ovariis nonnullis
inflorescentiae eiusdem, glomeruli sporarum faciliter in sporis dissoluti, sporae externae
10,5-14,5 um longae, atro- flavido- usque rufo-brunneae, in latere libero verrucis usque
0,5-1 um alltis.
ETYMOLOGY: named in the honour of the family of the outstanding Australian
mycologist, Dr. Roger G. Shivas, an excellent human being, a passionate collector of
various groups of plant parasitic microfungi, especially smut fungi, who discovered
and described numerous new species and genera, co-author of the comprehensive
monograph of the Fungi of Australia. The Smut Fungi (2008). Several interesting, rare
and even new species have been found by Roger’s wife, Marjan D.E. Shivas, and their two
daughters, Anthea and Gillian, who often accompany him on collecting trips to remote
parts of Australia and overseas.
146 ... Vanky
Fig. 10. Sori of Sporisorium shivasiorum in
the ovaries of Eulalia trispicata (type).
Habit. Bar = 1 cm.
10
Sort (Fig. 10) in all ovaries of an inflorescence, long cylindrical-fusiform,
0.5-1 x 5-15 mm, bearing on their acute tip remnants of two styles with
stigma, partly hidden by the floral envelopes, at first covered by a thick,
yellowish-brown peridium that later ruptures longitudinally in several places,
Ustilaginomycetes — 28 ... 147
disclosing the blackish-brown, granular powdery mass of spore balls and a
long, narrowing, often flattened central columella. SPORE BALLs (Figs. 13, 14)
subpolyhedrally irregular, rarely subglobose, ovoid or ellipsoidal, 30-110 x
40-—150(-180) ttm, reddish-brown, composed of tens or hundreds of spores that
separate by pressure. Sporgs (Figs. 13, 14) dimorphic, outer spores subglobose,
ellipsoidal to subpolyhedrally slightly irregular, 8-11.5 x 9-12 um, medium
dark yellowish-brown; wall uneven, 0.5-1(-1.5) um thick, thickest at the angles
and on the free surface, where it is evidently, densely, irregularly verrucose-
echinulate, warts up to 0.4 um high, contact sides apparently smooth to finely
punctate, spore profile smooth, on the free surface finely undulate. Inner spores
about the size of the outer spores, globoid to subpolyhedrally slightly irregular,
pale yellowish-brown; wall even, c. 0.5 um thick, apparently smooth to finely,
densely punctate. STERILE CELLS absent.
On Poaceae: Eulalia trispicata (Schult.) Henrard; SE Asia (Thailand). Known only from
the type collection.
Key to the smut fungi (Sporisorium) of Eulalia and Eulaliopsis
me opOre-Dallscephemeralasterile.cellstpresentempeneet-etat aa, sehen salen re ae), eter 2
- >poreballsssemr permanent ssterileicellsabsents eewere ts ter ee ee ee 5
Piaee opores\o 521355 long... eee ee eee Re fete ee ee S. pollinianum
- Spores lessthan.9t5 tim lon gees ate er eee: Meets Sere t wi. a P 3
omer soothe whole inilorescence, On edialiopsisima: Sues ene S. indicum
- SoumanitheispikeletskOnien aliamerer eee nee 2) Seed wae). ek. Ae, 4
4. Spores 5.5-9 um long; wall uneven, 0.5-1.5 um thick, in SEM densely
WeCCUCOSe, Wakts Ofthe samenkindine arene cert a ee eee oe S. guangxiense
- Spores 7-9.5 um long; wall even, 0.5 um thick, in SEM moderately densely
echinulate, between the spines minutely verruculose................. S. rarum
5. Spore balls 25-55 um long, of few spores (10-402); spores 8-14 tum long
EE AUG Sr ete ed. Ne ce eS ORS: S. polliniae
- Sporeiballs larseroiamore spores, spores larcer ae mre, © eee reresene eta tae 6
6. Outer spores up to 20 um long, spore wall 1-3 um thick........... S. crypticum
- Outeuspores upto lu Lim. long,.spore wall thinnetermem too ee ee 7
Z. SOtianispikelets (all) spore wallsle-2 (2.5) (in thicken eer rae S. eulaliae
~ Sori in ovaries (some or all of an inflorescence); spore wall 0.5-1.5 um thick ...8
8. Sori in some ovaries of an inflorescence; spore balls easily separating into spores;
outer spores 10.5-14.5 um long, with 0.5-1 um high warts........ S. trispicatae
~ Sori in all ovaries of an inflorescence; spore balls permanent; outer spores
V=12 iimmlong, with up to.0.4 tim high Warts). e). cou. a. S. shivasiorum
A new Thecaphora species on Arnica pumila (Asteraceae)
Thecaphora californica and T: cuneata are known from the USA on
Grindelia spp. A smut fungus, collected on Arnica pumila, was also attributed
148 ...Vanky
to T. californica by Fischer (1953:154). A study of the specimen shows that it is
specifically distinct and it is described as:
Thecaphora arnicae Vanky, sp. nov.
MycoBank: 511645
Typus in matrice Arnica pumila, USA, Colorado, Boulder, 19.VI.1919, leg. E. Bethel (ut
Thecaphora sp.). Holotypus WSP 34598!
Thecaphora arnicae distincta T. californica (Harkn.) G.P. Clinton, J. Mycol. 8:146, 1902
(typus in matrice Grindelia robusta, USA) ei proxime relativae speciei glomerulis sporarum
majoribus (17-45(-50) um longis) et (1-)3-25 sporis compositis, sporis majoribus. (12-15
(-17) x 13-20(-24) ym, radialiter 10,5-20 um longis, et ad superficiem liberam pariete
crassiore (2,5-3,5 um). In Thecaphora californica glomeruli sporarum 22-40 um longi,
e 6-20 (vel pluribus?) sporis compositi, sporae 9,5-13,5 x 9,5-15(-17) um, radialiter
9,5-17,5 um longae, pariete ad superficiem liberam 1,5-2,5 um crasso.
Sort in the flower heads. SporE BALLS (Figs. 15, 16) globose, subglobose, ovoid,
ellipsoidal, 15-45 x 17-45(-50) um, pale yellowish-brown, composed of (1-)
3-25 firmly united spores. Spores (Figs. 15, 16) subcuneiform, in surface view
rounded subpolygonally irregular, more rarely ovoid or elongated, 12-15(-17)
x 13-20(-24) um, radially 10.5-20 um long, yellow to pale yellowish-brown; —
wall of contact sides c. 0.5 um thick, smooth, on the free surface 2.5-3.5 um
thick, provided with irregular spiny warts, up to 1.5 um high, isolated or usually
confluent at their base into an irregular pattern.
On Asteraceae: Arnica pumila Rydb.; N America. Known only from the type collection.
Thecaphora arnicae differs from the closely related T. californica (Harkn.) G.P.
Clinton (1902:146; type on Grindelia robusta Nutt., USA), in having larger
spore balls (17-45(-50) um long), composed of (1—)3-25 spores, larger spores
(12-15(-17) x 13-20(-24) um, radially 10.5-20 um long), and thicker wall of
the free surface (2.5-3.5 um). In T. californica the spore balls are 22-40 um long,
composed of 6—-20(or more?) spores, the spores are 9.5-13.5 x 9.5-15(-17)
uum, radially 9.5-17.5 um long, the wall of the free surface is 1.5-2.5 um thick.
T. arnicae differs also from T. cuneata, in which the spore balls are 35-60(-70)
um long, composed of 15 to several tens of firmly united spores, which radially
are 12-26 um long, with a 2.5-4(-5) um thick spore wall on the free surface.
A new Ustilago species on Milium vernale (Poaceae)
Milium L. is a grass genus of 4 species in the tribe Stipeae of the subfam.
Pooideae (Clayton & Renvoize 1986:85). On Milium several smut fungi have
been described representing two species: Tilletia milii-vernalis Liro (Liro
1939:113; type on M. vernale, Turkey), and Ustilago milii (Fuckel) Liro (Liro
1924:78; = U. striiformis; type on M. effusum L., Germany). A different species,
with peculiar sori was found recently in the Herb. B (Berlin), which is described
as:
Ustilago milii-vernalis Vanky & H. Scholz, sp. nov.
MycoBaNnk: 511646
Ustilaginomycetes — 28 ... 149
Fig. 11. Sori of Ustilago milii-vernalis in the spikelets and floral pedicels of Milium vernale (type).
Habit, and enlarged a healthy spikelet, an infected spikelet and a group of four infected spikelets
in which the sori comprise also the floral pedicels.
Bars = 1 cm for habit, and 2 mm for detail drawing.
150 ...Vanky
Typus in matrice Milium vernale, Graecia, Makedonia, Florina, NW oppid. Ammohori,
40°46'24” N, °21°30°0.5” E, alt. 615 m.s.m., 10.V.2004, leg. R. et E. Willing 126494.
Holotypus in HUV 21458, isotypus in B.
Sori in spiculis tumores cinerei, bullati ad basin involucrorum floralium omnium, saepe
etiam ad pedicellos floralios, raro etiam fere ad involucra omnia, intacte apicem illorum
tantum relinquentes, prius epidermide tenuo cooperti, quo irregulariter rupto massam
brunneam, pulveream sporarum ostendentes. Vulgo spiculi omnes rami inflorescentiae
eiusdem affecti, cum ramis salvis permiscenter. Tam non raro spiculi pauci eiusdem rami
inflorescentiae tantum affecti. Sporae globosae, subglobosae, ovoideae vel ellipsoideae, raro
parum irregulares, 6,5-8 x 6,5-9,5(-11) um, pallide flavido-brunneae; pariete inaequali,
cca. 0,4 um in latere uno, cca. 0,8 um in parte dimidia opposita, qua spora etiam atrior,
superficies eius leniter, dense punctata-verruculosa, imago obliqua sporae in latere atriore
levis, in latere pallidiore leniter, dense undulata.
Sort (Fig. 11) in the spikelets as grey, bullate swellings on the base of all
floral envelopes, often also on the floral pedicels, rarely comprising nearly the
whole floral envelopes leaving intact only their tip, at first covered by the thin
epidermis that ruptures irregularly disclosing the brown, powdery mass of
spores. Commonly all spikelets of an inflorescence branch are affected, which
are mixed with healthy branches. However, not rarely only a few spikelets of
a certain inflorescence branch are infected. Spores (Figs. 17, 18) globose,
subglobose, ovoid or ellipsoidal, rarely slightly irregular, 6.5-8 x 6.5-9.5(-11)
tum, pale yellowish-brown; wall uneven, c. 0.4 um on one side, c. 0.8 um on the
opposite half, where the spore is also darker, surface finely, densely punctate-
verruculose, spore profile smooth on the darker side, finely, densely wavy on
the paler side.
On Poaceae: Milium vernale M. Bieb., S Europe. Known only from the type collection.
Ustilago milii-vernalis has unique morphology and distribution of the sori.
These characteristics, in combination with the spore morphology, differentiate
it from all known Ustilago species on Poaceae.
A new Ustilago species on Sclerachne punctata (Poaceae)
Sclerachne R. Br. is a genus with only one known species, S. punctata, in
the subtribe Chionachninae of the tribe Andropogoneae (Clayton & Renvoize
1986:371). No smut fungus is known on Sclerachne. A meagre specimen is
preserved in BPI, under the herbarium name Sphacelotheca sclerachnes Wakef.,
which is described as:
Ustilago sclerachnes Wakef. ex Vanky, sp. nov.
MycoBANnk: 511647
Sphacelotheca sclerachnes Wakef., nom. herb. — Typus in matrice Sclerachne punctata,
Thailand, Bangkok, 10.X1.1929, no further data. Holotypus BPI 195063! (a single
sorus).
Sori in ovariis, cca. 2 x 4mm, anguste ovoidei, inter involucra floralia distantes conspicui,
primum peridio pallide brunneo cooperti, quo irregulariter rupto massam pulveream
Ustilaginomycetes — 28... 151
fusco-brunneam sporarum ostendentes. Sporae globosae vel subglobosae, raro late
ellipsoidales, 5-6,5 x 5,5-6,5 um, mediocriter atro-flavidobrunneae; pariete aequali, cca.
0,5 um crasso, valde leniter, moderate dense punctato usque punctato-verruculoso, imago
obliqua sporarum levis. Cellulae steriles absentes.
Sori in ovaries (Fig. 12), c. 2 x 4 mm, narrow ovoid, showing between the
spreading floral envelopes, at first covered by a pale brown peridium that
ruptures irregularly disclosing the blackish-brown, powdery mass of spores.
SPORES (Figs. 19, 20) globose or subglobose, rarely broadly ellipsoidal, 5-6.5
x 5.5-6.5 um, medium dark yellowish-brown; wall even, c. 0.5 um thick, very
finely, moderately densely punctate to punctate-verruculose, spore profile
smooth. STERILE CELLS absent.
On Poaceae: Sclerachne punctata R. Br. (Chionachne punctata
(R. Br.) Jannink; Polytoca punctata (R.Br.) Stapf); SE Asia.
Known only from the type collection.
Ustilago sclerachnes differs from all known smut fungi on
members of the closely related Chionachne and Polytoca, as
shown in the key below.
Fig. 12. A sorus of Ustilago sclerachnes in the ovary of Sclerachne punctata Wy
(type). Bare mam: 12
Key to the smut fungi of Chionachne, Polytoca and Sclerachne
SMES OTL U1 LCR VAL CSM siete ee ane ee Peete cers Sa arehy Ge Ue. PRR ke 2
- Solute spi kelets-Omim the WnOleiMlOorescencety, 4. rye want eather 5
PM OD OLES O72 = DOM LINIONG # cee vegan oe ae eR ear ee tear Tilletia puneana
- SY eXAS RAS LONNIE Sic. nes RE AS Oe ses A oe ee a RA ear RR Ee De 3
SS DOlES2 550111 |ONG Meer, Beene... ee eernnere eres Tilletia chionachnes
- SPOresrsilallclan wm: 7 en en eee ete es Np ate eee © 4
peer SDOres 10 -24(—26) timlon Pes ees eee ee ey Tilletia kimberleyensis
- SPOTeS 5.50%: ION Gee ee ee eee ee ee Ustilago sclerachnes
5: Sori in the whole inflorescence; columella much branched................... 6
- Sot MLhespikelets/columellanoumuch branched eee ree ae er eee 7
6. Spores 8-10.5 um long, prominently echinulate ............ Sporisorium simile
- Spores 10.5-14.5 um long, densely, minutely echinulate Sporisorium reilianum
eae > POres O2o— lL vimlone: me eee a ee Sporisorium polytocae-barbatae
- Spores lat Gere rc yee mete rete eam res he Gt ea een eae nn 8
8. Spores 11-15 um long, mostly subpolyhedrally irregular, finely punctate-
echinulate, spore profile smooth to finely serrulate....... Sporisorium polytocae
> Spores 9.5-16(-19) um long, mostly rounded, evidently echinulate;
SDOLe DILOMICiscl alc een er ee te eet. Ce Sporisorium polytocae-digitatae
152 ... Vanky
Ustilaginomycetes — 28 ... 153
Figs. 19, 20. Spores of Ustilago sclerachnes on Sclerachne punctata in LM and in SEM (type).
Bars = 10 um.
Figs. 21, 22. Sectioned spore balls of Doassansiopsis guangdongensis on Sagittaria sp. in LM (type).
Fig. 21. Hand-cut, not stained. Fig. 22. Embedded in resin, thin-sectioned (0.7 tm) and stained
with new fuchsin and crystal violet (as described in Vanky et al. 2008, 172). hc = host plant cells,
cstc = cortical sterile cells, sp = spores, pstc = parenchymatous sterile cells.
Bars = 10 um.
Figs. 13, 14. Spore balls and spores of Sporisorium shivasiorum on Eulalia trispicata in LM and in
SEM (type).
Figs. 15, 16. Spore balls and spores of Thecaphora arnicae on Arnica pumila in LM and in SEM
(type).
Figs. 17, 18. Spores of Ustilago milii-vernalis on Milium vernale in LM and in SEM (type).
Bars = 10 um.
154 ...Vanky
New combinations
Thecaphora bulbinellae is a Urocystis (Urocystidaceae)
Talbot (1958:111) described a smut fungus on Bulbinella setosa (Liliaceae s.
lat.), under the name Thecaphora bulbinellae. It produces large, permanent, dark
reddish-brown spore balls in the flowers. According to Talbot, the spore balls
are composed of 20 to numerous spores. As all 58 known species of Thecaphora
infect dicotyledonous host plants, T: bulbinellae was investigated. Spore balls
bleached with hydrogen peroxide (H,O,) revealed that they were composed
of numerous spores surrounded by a single layer of dark coloured sterile cells.
Spore germination is not known and DNA analysis from old specimens was not
possible. Ultrastructural analyses revealed haustoria and simple septal pores
with membraneous caps and bands, typical, i.a., for the Urocystidaceae Begerow
et al. The genus Urocystis Rabenh. ex Fuckel is a large genus with 170 known
species parasitising both monocotyledonous and eudicotyledonous host plants
in 29 families. The sterile cells in the spore balls of Urocystis species are usually
hyaline or pale yellowish-brown. Dark, reddish-brown sterile cells in Urocystis
are rare (e.g. U. junci Lagerh.). Based on the results, T: bulbinellae is transferred
into the genus Urocystis as:
Urocystis bulbinellae (P.H.B. Talbot) Vanky, M. Lutz, R. Bauer & Piatek, comb. nov.
MycoBAank: 511648
BastonyM: Thecaphora bulbinellae P.H.B. Talbot, Bothalia 7:111, 1958. — Type on
Bulbinella setosa (= B. nutans), South Africa, Pretoria, Struben’s Valley, 24.V1.1957,
J. Toxopeus. Holotype PREM 41745, isotype HUV 21442!
Sori in all flowers of an inflorescence replacing the seeds with a black,
granular-powdery mass of spore balls, enclosed by the floral envelopes. SPORE
BALLS globose, subglobose, ellipsoidal to slightly irregular, 25-40 x 25-50 um,
yellowish to dark reddish-brown, composed of 20 to several tens of firmly
agglutinated spores surrounded by a layer of darker, larger, thick-walled sterile
cells, collapsed in dry specimens. SporEs varying in shape and size, globose,
subglobose, ovoid, usually subpolyhedrally irregular, 4-8 x 5-12 um, pale
yellowish-brown; wall even, c. 0.5 um thick, smooth. STERILE CELLS varying
in shape and size, globose, ellipsoidal, ovoid to elongated or slightly irregular,
5-12 x 5.5-15 um, medium dark reddish-brown; wall uneven, c. 0.5-2.5(-3)
tum thick, smooth on both contact and free surfaces. SPORE GERMINATION not
known.
On Asphodelaceae (Liliaceae s. |.): Bulbinella nutans (Thunb.) T. Durand & Schinz (B.
robusta Kunth; B. setosa (Willd. ex Schult.f.) T. Durand & Schinz; Anthericum setosum
Willd. ex Schult.f.); S Africa, two localities.
Urocystis bulbinellae deviates from typical Urocystis in having relatively small,
thin-walled, pale coloured spores of variable shape and size. Furthermore,
Ustilaginomycetes — 28... 155
it differs in having dark, thick-walled sterile cells. Spore ball building is also
different.
The generic position of Schizonella isolepidis
by KALMAN VANKy & Matruias Lutz
A peculiar smut fungus was described on the stems of Isolepis nodosa (= Ficinia
nodosa; Cyperaceae) from New Zealand, under the name Schizonella isolepidis
Vanky (Vanky & McKenzie 1990:251). The authors also discussed this fungus
and showed that S. isolepidis differs in several features from the other three,
rather uniform species of Schizonella. They concluded that its generic position
was uncertain.
To determine the phylogenetic position of Schizonella isolepidis the ITS and
LSU rDNA sequences of the type specimen of S. isolepidis were compared with a
dataset containing representatives of the Ustilaginales, including all Schizonella
and Tolyposporium species that had ITS and LSU sequences on GenBank.
Additionally we sequenced and analysed DNA of a specimen of Tolyposporium
neillii (G. Cunn.) Vanky & McKenzie (on Ficinia nodosa, New Zealand, Stewart
Island, Mason Bay, alt. c. 10 m, 14.11.1998, leg. E.H.C. McKenzie, C. & K.
Vanky, HUV 18533). For methods of DNA extraction, PCR and sequencing see
Lutz et al. (2004), for methods of phylogenetic analyses see Vanky et al. (2008).
GenBank accession numbers are given in Fig. 23.
The results of the molecular analyses are depicted in Fig. 23. Schizonella
isolepidis clusters as sister taxon of Tolyposporium neillii within a group
containing all Tolyposporium species sampled. In addition, Schizonella isolepidis
is well separated from Schizonella species. The morphological differences of
Schizonella isolepidis and Tolyposporium neillii, both growing on the same host
plant, correspond to considerable differences in the ITS and LSU nucleotide
sequences (19 + 7 base pairs differences between the two species). Consequently,
the following transfer is proposed:
Tolyposporium isolepidis (Vanky) Vanky & M. Lutz, comb. nov.
MycoBank: 511649
BasionyM: Schizonella isolepidis Vanky, in Vanky & McKenzie, New Zealand J. Bot.
28:251, 1990. — Type on Isolepis nodosa (= Ficinia nodosa), New Zealand, North Island,
Bay of Plenty, Moutohora (Whale Island), 4.1.1986, leg. R-E. Beever, Holotype HUV
14720, isotype PDD 56371.
Sori as small, subepidermal pustules on stems, often confluent, up to 10-15
cm long and surrounding the stems, at first lead-coloured, later the epidermis
flakes away, exposing the black, granular, semi-powdery mass of spores and
spore balls. Spores often in pairs or in balls of up to 10 spores, but isolated
spores are also present. SPORE BALLS irregular, 13-20 x 18-45 um, dark
reddish-brown, opaque. Mycelial remnants often present around the spore balls,
156 ... Vanky
ae Schizonella melanogramma DQ832212/D0832210
109/99} SSchizonella sp. DQ832218/D0832216
100/94] bStegocintractia luzulae DQ875353/AJ236148
Farysia chardoniana AY344968/AF009859
Tolyposporium isolepidis @Schizonella isolepidis) EU248950/EU246949
82/-
100/100
Tolyposporium neillii EU246951/EU246952
Tolyposporium junci AY344994/AF009876
715
Tolyposporium piluliforme DQ875345/AF009871
Macalpinomyces eriachnes AY740037/AY740090
400/100 (~~ Moesziomyces bullatus AY740153/AF009868
Ustilago maydis AY345004/AF 453938
100/85
Sporisorium sorghi AY740021/AF 009872
Moreaua fimbristylidis DQ875350/DQ875367
——().()5 substitutions/site
Ustilaginomycetes — 28... 157
and spores. SPORES very irregular in shape and size, subglobose, elongated,
subpolyangular to usually irregular, often with a flattened side, 10-16 x 14-24
um, dark reddish-brown; wall unequally thickened, 1-5 um wide, sometimes
with protuberances and light-refractive areas at the angles, surface irregularly
reticulate to cerebriform, meshes often incomplete. In SEM surface irregularly
ornamented by a combination of large, often incomplete meshes and warts.
SPORE GERMINATION unknown.
On Cyperaceae: Ficinia nodosa (Rottb.) Goethg. et al. (Scirpus nodosus Rottb.; Isolepis
nodosa (Rottb.) R. Br.); New Zealand.
Ustilago duriusculae is U. striiformis
by KALMAN VANKyY & MarTruias Lutz
Guo (2006:162) published Ustilago duriusculae on Carex duriuscula subsp.
rigescens (Franch.) S.Yun Ling & Y.C. Tang, from China, Qinghai Prov., Minhe,
Langtang, alt. 2580 m, 9.VIII.2004, coll. L. Guo & W. Li 2698. Holotype HMAS
95452, isotype HUV 20951! Paratype HMAS 95453, isoparatype HUV 20952!
The fungus produces long striae on the leaves between the veins, at first covered
by the epidermis that ruptures disclosing the dark brown, powdery mass of
spores. Spores subglobose, ellipsoidal, ovoid to slightly irregular, 7.5-12 x
8-15 um, yellowish- to reddish-brown; spore wall even, c. 1 um thick, finely
echinulate.
Guo (2006:162) remarked that the fungus “is close to Ustilago striiformis
(Westend.) Niessl on Poaceae from which it differs in parasitising different host
family.’ Unfortunately, the infected plants are sterile. Attempts to obtain base
sequences from fungal DNA failed. However, the 5’-end of the nuclear large
subunit ribosomal DNA (LSU) of the host plant was sequenced. (for methods
of DNA extraction, PCR and sequencing see Lutz et al. (2004), (GenBank
accession number: EU246953). Blast searches (Altschul et al. 1997) revealed
closest similarity of the sequence to sequences of Poaceae. Consequently,
Ustilago duriusculae L. Guo is considered to be a synonym of U. striiformis
(Westend.) Niessl (syn. nov. by Vanky & M. Lutz) and it remains valid, that
Ustilago spp. are restricted to members of the Poaceae.
Fig. 23. Bayesian inference of phylogenetic relationships within the sampled Ustilaginales: Markov
chain Monte Carlo analysis of an alignment of ITS and LSU rDNA base sequences using the
GTR+I+G model of DNA substitution with gamma distributed substitution rates and estimation
of invariant sites, random starting trees and default starting parameters of the DNA substitution
model. A 60% Majority-rule consensus tree computed from 18 000 trees that were sampled
after the process had reached stationarity is shown. The topology was rooted with Moreaua
fimbristylidis. Numbers on branches before slashes are estimates for a posteriori probabilities,
numbers on branches after slashes are percentage bootstrap values of 1000 replicates. Branch
lengths were averaged over the sampled trees. They are scaled in terms of expected numbers of
nucleotide substitutions per site. (Prepared by M. Lutz).
158 ...Vanky
About the genus Melanotaenium (Melanotaeniaceae)
The genus Melanotaenium de Bary (Bary 1874:105), with its type
M. endogenum on Galium mollugo (Rubiaceae), was described for a Protomyces-
or Entyloma-like fungus. It has single, dark coloured spores embedded in the
host tissue. Successively, a large number of parasitic microfungi representing
a heterogeneous assemblage from many different host plants have been
transferred into or described in Melanotaenium. Based on morphological,
biological, ultrastructural and molecular characters, as well as on host plant
criteria, of the c. 60 Melanotaenium species remained eleven. These are:
1. M. adoxae (Bref.) S. Ito, type on Adoxa moschatellina L. (Adoxaceae), Germany,
2. M. antirrhini Vienn.-Bourg. ex Vanky, type on Antirrhinum latifolium
Mill. (Scrophulariaceae), France, 3. M. arnaudianum Cif., type on Helleborus
niger L. (Ranunculaceae), Italy, 4. M. cingens (Beck) Magnus, type on Linaria
genistifolia (L.) Mill. (Scrophulariaceae), Austria, 5. M. endogenum (Unger)
de Bary, type on Galium mollugo L. (Rubiaceae), Austria, 6. M. euphorbiae
(L.W. Lenz) M.D. Whitehead & Thirum., type on Euphorbia heterophylla L.
(Euphorbiaceae), USA, 7. M. gunnerae G.P. Clinton ex Vanky & R. Bauer, type
on Gunnera magellanica Lam. (Gunneraceae), Chile, 8. M. hypogaeum (Tul.
& C. Tul.) Schellenb., type on Linaria spuria (L.) Mill. (= Kickxia spuria (L.)
Dumort; Scrophulariaceae), France, 9. M. jaapii Magnus, type on Teucrium
montanum L. (Lamiaceae), Germany, 10. M. spermacoces ‘Thirum. et al., type
on Spermacoce hispida (Rubiaceae), India, and 11. M. tochinaianum S. Ito, type
on Peracarpa carnosa Hook. f. & Thomson (Campanulaceae), Japan.
With the exception of M. arnaudianum and M. spermacoces, species
typically have sori that are swollen, forming pustules, swellings, sometimes
considerable tumours, usually on the stems and also on the hypocotyls, petioles
and in the leaf veins or leaves, the spores are relatively large, up to 20, 24 or
even 28 um long, dark reddish-brown, with a two-layered, up to 2.5 or 4, rarely
5.5 um thick wall (comp. also Vanky 2002:98). In contrast, M. arnaudianum
and M. spermacoces form spots on the leaves, and the spores are small, 9-11,
respectively 7.5-12.5 um. The spore wall in M. arnaudianum is only 1-1.5 um
thick, and the spores are hyaline or brownish. According to Ciferri (1924a:58),
Arnaud found this fungus on Helleborus near Savoia and considered it to be
Entyloma ranunculi. Based on the description, and after comparison with the
other Melanotaenium species, I consider that the place of M. arnaudianum is
within the genus Entyloma as:
Entyloma arnaudianum (Cif.) Vanky, comb. nov.
MycoBank: 511650
BAsIONYM: Melanotaenium arnaudianum Cif., Bull. Soc. Bot. Ital. 1924:58, 1924. —
Type on Helleborus niger, Italy, Pausula, near Macerata, date? coll. R. Ciferri (type
where?).
Ustilaginomycetes — 28 ... 159
Sor! in leaves and petioles as elongate spots with indefinite margins, up to 5
cm long or longer by confluence, initially pale then reddish-brown or brownish
violet, covered by the epidermis. Spores intercellular, globose, 9-11 um in
diam., at first hyaline, later brownish, contents sometimes guttulate; wall
two-layered, 1-1.5 um thick, smooth. [No material was available for study.
Description taken from the original. ]
On Ranunculaceae: Helleborus niger L.; Europe (France, Italy).
Regarding Melanotaenium spermacoces (Thirumalachar et al. 1956:10), it
was described on Spermacoce hispida L., from India, Bombay, Ambernath,
15.VHI1.1954, coll. M.K. Patel, G.W. Dhande & V.V. Bhatt. (type where?, not
in HCIO, BPI, IMI). The original description is summary: “Sort forming dull
grey, diffuse, circular leaf spots without inciting malformation of the host.
SPORES scattered in the intercellular spaces of the mesophyll, deep olivaceous-
brown, 7.5-12.5 um in diam., with a small hyphal appendage attached to one
end. SPORE GERMINATION unknown.”
Furthermore, the authors remarked that the “Infection is confined mostly
to lower leaves”. “The spores are developed sparsely at the end of the hyphal
branches. A small remnant of the hyphae remains persistent appearing as a
small hyphal appendage”. “The spores are loosely grouped and very small in
size’. These characters exclude Melanotaenium spermacoces not only from the
genus Melanotaenium but also from the smut fungi (excluded here).
Key to the species of Melanotaenium, based on host taxonomy
On Adoxaceae: Adoxa; in stems, rhizomes and leaves...............-.0-- M. adoxae
mim ampantilaceae: Peracarpa; On stems. es. ee ae M. tochinaianum
Bern piorvidceae te Uv Ol ola sONsleIsteeee fr ee M. euphorbiae
On Gunneraceae: Gunnera; on petioles and leaf veins ................. M. gunnerae
On Lamiaceae: Galeopsis, Lamium, Teucrium,
SIPUNGeroTOUNGistemis andy POCOLVSm em We eae erates tras of) ee ee M. jaapii
On Rubiaceae: Asperula, Galium, on stems and leaves............... M. endogenum
On Scrophulariaceae:
1. On Antirrhinum; on stems; spores 14-20 um long; wall more or less evenly
Bicone ele, (LW ICO me tte ee ee wen See eee M. antirrhini
5 INOUONPANUILHINUM eDOtessLO— 24. LIMON 2. ee yer Gee ane eee eee p}
2. On Kickxia; on hypocotyl as large, cylindrical swelling ......... M. hypogaeum
= One idria- on stems and leaves as PUSLULCS. 1 aes a tee ere eee M. cingens
Tuburcinia simplex is a good species in Urocystis (Urocystidaceae)
On species of Anemone (Ranunculaceae) several Urocystis species have been
described. The following eight species could be recognised: 1. U. anemones
160 ...Vanky
(Pers. : Pers.) G. Winter (Winter 1880:160; type on A. nemorosa L., Germany),
2. U. antipolitana Magnus (Magnus 1879:214; type on A. coronaria L., France),
3. U. antucensis (Liro) Piatek (Piatek 2007:93; type on A. antucensis Poepp.,
Chile), 4. U. japonica (Henn.) L. Ling (1945:3; type on A. japonica Siebold &
Zucc., Japan), 5. U. novae-zelandiae (G. Cunn.) G. Cunn. (1945:344; type on
A. tenuicaulis (Cheeseman) Parkin & Sledge, New Zealand), 6. U. subnuda
(Liro) Zundel (Zundel 1953:334; type on A. cylindrica A. Gray, USA),
7. U. pseudoanemones Denchev et al. (Denchev et al. 2000:453; type on
A. flaccida F. Schmidt, Japan), and 8. U. sinensis L. Guo (Guo 2005:270; type on
A. rivularis Buch.-Ham. ex DC., China).
An additional smut fungus, described by Liro as Tuburcinia simplex on Anemone
caroliniana, belongs to the genus Urocystis as:
Urocystis simplex (Liro) Vanky, comb. nov.
MycoBank: 511651
BastonyM: Tuburcinia simplex Liro, Ann. Univ. Fenn. Abo., Ser. A, 1:59, 1922. — Type
on Anemone caroliniana, USA, Texas, Austin, 13.11.1901, W.H. Long, Holotype in
H!, isotype BPI 181309.
Sori on leaves forming pustules up to 1 cm in diam., at first covered by the
epidermis that ruptures irregularly disclosing the black mass of spores and
spore balls. SPORE BALLS composed of 1, rarely 2 spores, usually naked or
with a few sterile cells. SpoREs globose, ovoid, ellipsoidal or slightly irregular,
11-15 x 13-19 um, reddish-brown; wall 0.8-1.2 um thick, apparently smooth.
STERILE CELLS globoid to elongated, 10-19 um long, yellowish-brown; wall 1-3
um thick, smooth, collapsed in old specimen.
On Ranunculaceae: Anemone caroliniana Walter; N America (USA).
Typical for Urocystis simplex are that the spore balls are restricted to mostly a
single spore, and sterile cells are often absent.
Key to the Urocystis species of Anemone and Anemonella
1... Spote balls surrounded by,a’ continuous layer of sterile.cells. 3.25) oe p)
- Spore balls surrounded partly by a discontinuous, partly by a continuous layer of
sterile:cells orsterile cells lackino: 7. sae, cet Oe ae ee ee eee 5s
22,4, spore balls composedior 1-20(-30)' spores’ 4. U. novae-zelandiae
- Spore balls composed of | l2)spotes: = 2 a. ate oe eae eee U. japonica
3), spore balls lackine sterile cells or, with:a fewsstemle cells" 5... et ae ee +
- Spore balls surrounded by an incomplete or complete layer of sterile cells...... 5
Ay & opore' balls‘composed O61 (2) spores= asst ee ae tenn anes U. simplex
- Spore balls composed of 1-4(=5) sporest sun eh este, oe eae aoe U. subnuda
5. Spore balls composed of (0—)1-2(-3) spores; sterile cells few or lacking
ee ee SOR i ee Se ee Ee U. anemones
Ustilaginomycetes — 28 ... 161
on ropore ballsccomposed of 1=4(E6)’sporesee. 9.08. U. antucensis
- SPO reDauis cOMIDOSEWIOL Ol eSDOLCSEy ere ms oe eee 6 ok cs Satay ee 7
fee pore balls composed of 1-6(—7) spores... 2.22... ..2....0.. U. antipolitana
- BpOle balls cCOMpOsed. Orley (alt ECDOTCQe ment ttrs fy cin. rue ciate a eee ene ee 8
ga) pore balls 16—40 pm long; spores 9:5-17 tim long..-.-............ U. sinensis
- Spore balls 20-70 um long; spores 13-21(-24) um long..... U. pseudoanemones
New members of the genus Gjaerumia (Urocystidaceae)
The host plants of c. 180 known species of Entyloma are restricted to
dicotyledonous host plants in 25 families (Vanky 2002:56). Two “Entyloma”
species on Liliaceae s. |., E. eremuri and E. muscari, do not belong to this
genus. A third species, E. ossifragi Rostr., on Narthecium ossifragum (L.) Huds.,
was found to represent a new genus, Gjaerumia R. Bauer et al. (Bauer et al.
2005:109). The other two “Entyloma” species on Liliaceae s. |. are transferred
into Gjaerumia as:
Gjaerumia eremuri (Schwarzman) Vanky, comb. nov.
MycoBank: 511652
BasionyM: Entyloma eremuri Schwarzman, FI. spor. rast. Kazakhstana 2:266, 1960. —
Type on Eremurus tianschanicus, Kazakhstan, Alma-Ata Region, surroundings of
Alma-Ata, south slope, 18.VI.1946, S.R. Schwarzman, AA (n.v.); South Kazakhstan
Region, Talasskij Alatau, road towards Djabagly, 8.VI.1951, A.N. Ustimenko, AA
(n.v.). (No type designated).
Sori forming rounded-elongate, obscure, violaceous leaf spots. SPoREs few,
embedded in the mesophyll and leaf epidermis, globose, angularly globose or
ellipsoidal, 5-14 x 5-16 um, mostly 7-9 um in diam.; young spore walls are
smooth or dotted, hyaline, 1.5-2 um thick, when mature smoky cinnamon
brown, 1-1.5 um thick. ANAMORPH absent.
On Asphodelaceae (Liliaceae s. |.): Eremurus tianschanicus Pazij & Vved. ex Pavlov; C
Asia.
Material not seen. Description taken from the original.
Gjaerumia muscari (Pass. ex J. Schrét.) Vanky, comb. nov.
MycoBank: 511657
Basionym: Entyloma muscari Pass. ex J. Schrot., in Cohn, Beitr. Biol. Pfl. 2:371, 1877,
as ‘Entyloma Muscari (Passerini). — Type on Muscari comosum, Italy, Parma, G.
Passerini, PARMA (n.v.).
Sort in leaves as elliptical or rhomboidal, c. 1 cm long, flat, brownish discoloured
spots. SPoRES in the host tissue between the parenchyma cells, globose, c. 11
ttm in diam., pale brown; wall thin, smooth.
On Hyacinthaceae (Liliaceae s. |.): Muscari comosum (L.) Mill.; Europe. Known only
from the type locality.
Material not seen. Description taken from Schréter (1877:371). According to
Schroter, E. muscari is morphologically close to E. corydalis. Ciferri (1924b:95)
162 ...Vanky
studied the type and was not completely convinced that it was an Entyloma
species.
Key to the species of Gjaerumia, based on host taxonomy
Lo, OnsNarthecium, Spores 10-14 5( 07 ) i One: ae aces ere G. ossifragi
- NOt On N@rtheciuin seta tae, cbt OF ee retain eae ee ee J:
2 ge OD LEPEMUPUS SSP OLeS 5 Og I) ODO were ean) en ye eee G. eremuri
- On. Muscari. Spores ¢. aul pina iainieteiae ie wea Patent te ewe G. muscari
An additional species of Bauerago (Microbotryaceae)
Seven species of Bauerago Vanky are known, parasitising host plants in
Commelinaceae, Cyperaceae and Juncaceae. An additional species, described as
Ustilago is transferred into this genus as:
Bauerago tinantiae (J.C. Lindg.) Vanky, comb. nov.
MycoBank: 511653
BasionyM: Ustilago tinantiae J.C. Lindgq., Revista Fac. Agron. Univ. Eva Peron, Ser. 3,
29:253, 1953. — Type on Tinantia fugax, Argentina, Salta Prov., Capital, Quebrada
de San Lorenzo, alt. 1500 m, 19.III.1952, leg. H. Sleumer. Holotype LPS 22062,
isotype HUV 13705!
SORI in ovaries. Spore mass chestnut-brown, powdery. Spores globose,
subglobose to ellipsoidal, 9.5-11 x 9.5-13 um, light brown; wall 1-1.5 um thick,
finely reticulate, 5-7 meshes per spore diam., areolae 1-1.5 um wide, muri
c. 0.8 um high, in median view blunt, 17-22 on the equatorial circumference, in
SEM muri and interspaces finely verruculose. SPORE GERMINATION (Lindquist
1953:254) results in four-celled basidia, producing laterally and terminally
ovoid basidiospores.
On Commelinaceae: Tinantia fugax Scheidw., Tinantia sp.; S America (Argentina).
Species of Ustilago are restricted to host plants in the Poaceae (see also chapter
U. duriusculae in this paper). The recombination of U. tinantiae into Bauerago
is based on spore germination, described and illustrated by Lindquist. The
ascomycetous ultrastructure, observed by R. Bauer (pers. comm.) probably
belongs to saprobic fungi often present in the sori of smut fungi collected in the
tropics and moulded.
Sorosporium solidaginis is a Thecaphora (Glomosporiaceae)
Schofield (Bessey, Contr. Bot. Dep. Univ. Nebr. 3:48, 1892; n.v.) described
Sorosporium cuneatum, in the flowers of Grindelia squarrosa, from the USA. One
year later, Ellis & Everhart (1893:156) described Sorosporium solidaginis, in the
flowers of Solidago missouriensis, from the USA. Two years later, Ellis & Everh.
(1895:363) declared that S. solidaginis “is the same as S. cuneatum Schofield”.
However, a comparison of the types revealed that there are differences in the
Ustilaginomycetes — 28 ... 163
spore ball and spore morphology, which justifies treating them as two species,
both within the genus Thecaphora.
Thecaphora solidaginis (Ellis & Everh.) Vanky, comb. nov.
MycoBAnk: 511654
BASIONYM: Sorosporium solidaginis Ellis & Everh., Proc. Acad. Nat. Sci. Philadelphia
1893:156, 1893. — Type on Solidago missouriensis, USA, Cansas, Rooks Co.,
Rockport, 28.11.1892, E. Bartholomew 531. Lectotype (designated here) BPI
179719; isolectotypes BPI 179136, BPI 179718, HUV 20819!
Sori destroying all flowers in a head, which are deformed, composed of
remnants of host tissues and a reddish-brown, granular powdery mass of spore
balls. SpoRE BALLS globose, subglobose, ovoid, ellipsoidal to slightly irregular,
25-40(-45) x 25-45(-50) um, pale yellowish-brown, composed of 8 to several
tens of firmly united spores which rather break than separate by hard pressure.
SPORES mostly subcuneiform, in surface view rounded subpolyangularly
irregular, rarely elongate, 8-12 x 9.5-18 um, radially 8-18(-20) um long, pale
yellow to pale yellowish-brown; wall of contact sides c. 0.5 um thick, smooth,
wall of the free surface 1.5-2.5(-3) um thick, finely, moderately densely,
irregularly verrucose, warts up to 1.5 um high, isolated or 2(-3) fused, spore
profile finely, irregularly wavy to serrulate.
On Asteraceae: Solidago missouriensis Nutt; USA. Known only from the type
collection.
Thecaphora solidaginis differs from T. cuneata (Schofield) G.P. Clinton (type on
Grindelia squarrosa (Pursh) Dunal, USA), especially in the morphology of the
spore balls and spores. The spore balls in T. cuneata are 35-60(-70) um long,
composed of 15 to several tens of spores which radially measure 12-26 um;
wall of the free surface is 2.5-4(-5) um thick.
Ustilago stiparum is a Sporisorium (Ustilaginaceae)
Sporisorium stiparum (Speg.) Vanky, comb. nov.
MycoBANK: 511655
BastonyM: Ustilago stiparum Speg., Anales Mus. Nac. Buenos Aires, Ser. 3, 12:288,
1909. — Sorosporium stiparum (Speg.) Zundel, Mycologia 43:269, 1951. — Type on
Stipa sp., Argentina, Mendoza, Lujan de Cuyo, 1.1.1908, C. Spegazzini. Holotype
LPS 3003, isotypi Herb. Hirschhorn 3003, HUV 15925!
Sori destroying the whole inflorescence, long linear, 4 cm or longer, more or
less hidden by leaf sheath, at first covered by a thin, whitish peridium which
breaks up in long strips exposing the dark brown, granular-powdery mass of
spore balls intermixed with long columellae. SPORE BALLS subglobose, ovoid,
ellipsoidal, elongated or subpolyhedrally irregular, 30-80 x 40-90(-100) um,
dark reddish-brown to subopaque, composed of tens or hundred(?) of spores
that separate by pressure. SporEs globose, ovoid or subpolyhedrally irregular,
164 ... Vanky
6.5-9 x 7-9.5(-10.5) um, yellowish-brown; wall even, 0.5-0.8 ttm thick, finely,
densely punctate-verruculose, spore profile smooth to very finely undulate or
minutely serrulate. STERILE CELLS not seen.
On Poaceae: Stipa sp.; Argentina. Known only from the type locality.
The fragment of the isotype, obtained from the late Dr. Elisa Hirschhorn,
does not permit identification of the host plant and a description of the soral
characters, which are taken from the original and Zundel (1953:74).
Synonyms
Entyloma crepidis-tectorum is identical with E. zacintha
Piatek (2006:165) described Entyloma crepidis-tectorum on the leaves of
Crepis tectorum from Denmark. He found the spores measured (10-)13-18
(-20) x (13-)15-21(-22) um, and unevenly, 2-5(-7) um thick spore wall. I
checked the holotype and obtained smaller values, namely 9-14.5 x 11-17
(-18.5) um for the spores and 1-5(-5.5) um for the wall thickness. After I
discussed the problem with Dr. Piatek, I checked another sorus of the holotype
and obtained (9.5-)10-15(-16) x (10.5-)12-18(-19) um for the spores, and
up to 5 um for the spore wall thickness. Obviously, there is a certain variation
in spore morphology on the same host plant, using the same method. Because
the spore morphology of E. crepidis-tectorum is very close to that of Entyloma
zacintha (type on Crepis zacintha), we came to the conclusion that it is best to
treat these smuts as one species:
Entyloma zacintha Vanky, Mycotaxon 36:474, 1990.
Type on Crepis zacintha, Greece, Rhodos, between Agaia Isodoros and Laerma, alt. c.
200 m, 27.IV.1978, K. Vanky. Holotype HUV 7174!
Entyloma crepidis-tectorum (as “crepidis-tectori’) Piatek, Polish Bot. J. 51:165, 2006.
— Type on Crepis tectorum, Denmark, Yullandia, near Ryga, VII.1911, C. Ferdinandsen,
BPI 174632! (syn. nov.).
Sort in leaves as rounded, thin, amphigenous, scattered or gregarious spots,
0.5-2 mm in diam., at first whitish, then yellow and finally light brown. Spores
solitary or usually densely crowded in small groups, globose, ovoid or more
or less irregular by mutual pressure, 10-15 x 13-19 um, subhyaline to pale
yellowish-brown; wall two-layered, the inner layer c. 0.5 um thick, the outer
layer often unevenly thickened, 1-4(-6) um wide, smooth. ANAMORPH not
seen.
On Asteraceae: Crepis tectorum L., C. zacintha (L.) Loisel. (Lapsana zacintha L.; Zacintha
verrucosa Gaertn.); Europe.
Entyloma lavrovianum is E. hieracii
Schwarzman (1960:297) described Entyloma lavrovianum on the leaves of
several Hieracium species. Comparison of the syntypes with the lectotype of
Ustilaginomycetes — 28 ... 165
E. hieracii did not reveal any essential difference in spore morphology. Therefore
they are considered synonyms as:
Entyloma hieracii Syd. & P. Syd. ex Cif. (Ciferri, Bull. Soc. Bot. Ital. 1924:50, 1924a).
Entyloma hieracii Syd. & P. Syd., Ann. Mycol. 16:244, 1918; nom. nud.. — Lectotype
(design. by Vanky 1985:69) on Hieracium murorum L., Germany, Berlin, at Schlachtensee,
VI1.1897, P. Sydow, HUV 1174!; isolectotypes in Sydow, Ust. no. 143 (as “Entyloma
calendulae’).
Entyloma crepidis Kawecka-Starmachowa, Glownie i Sniecie Polski 2:173, 1939. — Type
on “Crepis praemorsa (L.) Tausch” (= misnamed Hieracium sp., det. Vanky 1990:474),
Ukraine, Kolomyya (formerly Poland, Kolomya), Bania Berezowska, Rokieta Mt.,
V1.1913, A. Wroblewski, KRAM-F 2521! (syn. by Vanky 1990:474).
Entyloma lavrovianum Schwarzman, Fl. spor. rast. Kazakhstana 2:297, 1960. — Syntypes
on Hieracium ganeschinii Zahn, and H. korshinskyi Zahn, both from Kazakhstan, Alma-
Ata Region, Zailijskij Alatau, Great Alma-Ata Gorge, 19.VIII.1958, S.R. Schwarzman,
AA; isosyntypes HUV 12112! & HUV 12113! (syn. nov.).
For a description see Vanky (1994:95-96).
Urocystis rechingeri and U. radicicola represent the same species
A study of the types of Urocystis rechingeri and U. radicicola revealed that
they are identical. Both species have been collected only once, no doubt because
of their hidden sori on the roots of Ranunculus species.
Urocystis rechingeri Petr., Sydowia 20:278, 1968 (‘1966’).
Type on Ranunculus sp. (= R. komarovii, det. K.H. Rechinger), Afghanistan, Kabul,
below Istalif, alt. 1850 m, 1.V.1967, K.H. Rechinger. Holotype W 9187!
Urocystis radicicola H. Scholz & I. Scholz, Nova Hedwigia 47:69, 1988. — Urocystis
anemones f. radicicola Thomas (nom. herb.). — Type on Ranunculus arvensis, Germany,
Thiiringen, Ohrdruf, 8.V1.1893, F. Thomas. Holotype HBG! (syn. nov.).
Sori as large cylindrical, fusiform or curved swellings on the upper part of the
root or on the hypocotyl, 4-8 mm thick, up to 12 mm long, covered by host
tissue. They have an irregularly, incompletely polycystic structure enclosing
the black, semiagglutinated mass of spore balls which are liberated when the
host tissues are decomposed. SPORE BALLS globose, ellipsoidal, elongated to
irregular, 18-35 x 20-50 um, reddish-brown, composed of (0—)1-4(-5) spores
and an incompletely, rarely completely surrounding layer of sterile cells. SPORES
globose, ellipsoidal or irregular, with one or several flattened sides, 11-16 x
13-19 um, reddish-brown; wall even, 0.5-0.8 um thick, apparently smooth to
very finely punctate-verruculose. STERILE CELLS variable in shape and size,
subglobose, ellipsoidal, elongated or irregular with slightly flattened contact
sides, 5-12 x 6-15 um, pale yellowish-brown; wall uneven, 0.5-2 um thick,
thin on the free surface, thick on the contact sides, smooth.
On Ranunculaceae: Ranunculus arvensis L., R. komarovii Freyn; Europe (Germany),
Asia (Afghanistan).
166 ...Vanky
In the protologue of Urocystis rechingeri, the spore measurements were given
as 6-11 x 10-16 um. Under standard conditions, i.e. heated in lactophenol,
I obtained larger values, as given in the description above, identical with those
of U. radicicola. Because of the identical sori, spore morphology and host plant
genus, these two names are considered synonyms.
Ustilago moehringiae is a synonym of Microbotryum duriaeanum
Togashi & Maki (1940:139) described Ustilago moehringiae in the ovaries
of Moehringia platysperma from Japan. It was subsequently placed into the
genera Microbotryum and Haradaea. A study of its type and comparison with
Microbotryum duriaeanum could not reveal any morphological differences.
Consequently, they are considered heterotypic (taxonomic) synonyms as:
Microbotryum duriaeanum (Tul. & C. Tul.) Vanky, Mycotaxon 67:43, 1998.
Ustilago duriaeana Tul. & C. Tul., Ann. Sci. Nat. Bot., Sér. 3, 7:105, 1847. — Haradaea
duriaeana (Tul. & C. Tul.) Denchev & H.D. Shin, in Denchev etal., Mycol. Balcan. 3:72,
2006. — Type on Cerastium glomeratum Thuill., Algeria, Tlemcen, 30.V.1842, R.C.
Durieu de Maisonneuve. Holotype in PC!
Ustilago moehringiae Togashi & Y. Maki, Ann. Phytopathol. Soc. Japan 10:139, 1940.
Microbotryum moehringiae (Togashi & Y. Maki) Vanky, Mycotaxon 67:46, 1998. —
Haradaea moehringiae (Togashi & Y. Maki) Denchev, in Denchev et al., Mycotaxon
98:182, 2007('2006’). — Type on Moehringia platysperma Maxim. (= M. trinervia var.
platysperma (Maxim.) Makino), Japan, Chikuzen Prov. (= Fukuoka Pref.), Yoshikawa-
mura, 7.V.1938, Y. Maki. Holotype in TNS, isotype in SAPA! (syn. nov.).
Sor! in ovules, filling the capsules with at first agglutinated, later powdery, dark
purplish-brown spore masses. Infection systemic, all capsules of an infected
plant being affected. Sporss globose, subglobose to ovoid, 11-15 x 12-17 um,
light to medium purplish-brown, reticulate, 4-8 meshes per spore diam., muri
1-2(-2.5) um high, 16-22 on the spore circumference; interspaces in SEM with
a smooth or slightly roughened tubercle.
On Caryophyllaceae (subfam. Alsinoideae): Arenaria, Cerastium and Moehringia species,
Europe, N & NE Africa, Asia, N America.
Lectotype
Entyloma lobeliae Farl., Bot. Gaz. (Crawfordsville) 8:275, 1883.
Lectotype on Lobelia inflata L., USA, Maine, Gilead, VHI.1882, W.G. Farlow, (designated
here) BPI 175796.
Sort on the leaves forming rounded or polyangular, flat, amphigenous spots,
1-10 mm in diam., first whitish, later pale yellowish-brown. Spores globose,
subglobose, ellipsoidal or slightly irregular, 8-12 x 9-14 um, subhyaline to pale
yellow; wall even, 1-1.5 um thick, smooth. ANAMORPH present.
On Lobeliaceae: Lobelia cliffortiana L., L. inflata L., L. laxiflora Kunth, L. purpurascens
R.Br., L. spicata Lam.; N & C America.
Ustilaginomycetes — 28 ... 167
Entyloma lobeliae was described by Farlow, collected on Lobelia inflata, from
USA, Maine, Gilead, on August 18882. No type is preserved in FH. A neotype
for it was designated by Piepenbring (2003:211) on “Lobelia sp., USA, N.H.,
Shelburne, 1883, FH”. An isotype of E. lobeliae is preserved in BPI 175796.
According to Article 9.17(a) of the ICBN Vienna, the isotype supersedes the
designated neotype, which is not mentioned in the protologue.
Doassansiopsis guangdongensis, an unusual species
Recently, in addition to the 13 known species of Doassansiopsis,
(Urocystidales, Doassansiopsidaceae) a new species, D. guangdongensis S.H. He
& L. Guo (He & Guo 2007:101) was described on Alisma sp. from China. The
genus Doassansiopsis is characterised by permanent spore balls, embedded in
the host tissue of aquatic or paludal plants. The spore balls are composed of
a central mass of parenchymatous sterile cells, surrounded by spores and an
outer cortical layer of sterile cells. There is another group of 36 similar smut
fungi, the “Doassansia” group, classified in 11 genera within the Doassansiales,
Doassansiaceae. These genera differ from each other by the composition and
arrangement of the spores, sterile cells and/or hyphae in the balls (comp.
Vanky 2002:9, Pl. I, Figs. 1-11, and Vanky & Shivas 2006:16, Figs. 3-6). The
composition and structure of the spore balls can be studied best in embedded,
sectioned and stained balls, because the poorly pigmented and often very
similar spores and sterile cells are difficult to recognise and differentiate in
hand-cut, not properly stained balls. Study of the isotype of D. guangdongensis
revealed another structure of the spore balls than given in the protologue. In
the original description, the pigmented, outer layer of sterile, cortical cells were
mistaken for spores, host plant cells surrounding the balls as sterile cells, and
the spores as a part of the central mass of sterile cells. A revised description of
this smut is:
Doassansiopsis guangdongensis S.H. He & L. Guo, Mycotaxon 101:2, 2007,
emend. Vanky
Type on Sagittaria sp., China, Guangdong (no further data). Holotype HMAS 17625,
isotype HUV 21445!
Sori on the leaves forming yellowish brown, flat spots, from 1-2 to 10 mm
in diameter or larger by confluence, with spore balls embedded in the host
tissue as densely situated, pale brown dots. SPORE BALLS (Figs. 21, 22) globose,
subglobose, ellipsoidal, elongated or slightly irregular, with flattened sides,
c. 80-200 x 100-320 um, pale olivaceous brown, composed of a central mass
of empty, parenchymatous, sterile cells, surrounded by 1-3(-4), irregular,
often interrupted layers of spores, and a thick, well-developed cortical layer
of empty, thick-walled, pigmented sterile cells. SpoREs subglobose, ellipsoidal
or rounded irregular with slightly flattened contact sides, 5-9 x 5.5-10.5
168 ...Vanky
um, pale olivaceous brown; wall even, 0.5-0.7 um thick, smooth, content
homogenous. STERILE CELLS globoid or elongated, usually subpolyhedrally
or polyhedrally irregular, 5-12.5 x 5.5-20 um, pale olivaceous brown tinted;
wall even, 0.2-0.3 um thick. CORTICAL CELLS subglobose, ellipsoidal, rounded
irregular with slightly flattened contact sides, tangentially or radially elongated
or isodiametric, (5-)6-10.5 x (5-)7-13.5(-16) um, medium dark olivaceous
brown; wall uneven, 0.7-1.5 um thick, smooth.
On Alismataceae: Sagittaria sp.; E Asia. Known only from the type collection.
D. guangdongensis is closest to D. intermedia (Setch.) Vanky (type on Sagittaria
latifolia Willd., USA), in which there are 3-6 compact layers of spores in the
balls.
A tentative key to the genera of smut fungi
(Ustilaginomycotina p.p., and Microbotryales)
based on selected characters and on host plant taxonomy
During the last few decades numerous genera of smut fungi have been .
described, many as a result of the application of ultrastructural and molecular
phylogenetic methods. The Illustrated Genera of Smut Fungi (Vanky 2002)
listed 77 genera. Today 91 genera are known. and a key is given to these genera
below. For detailed descriptions and illustrations see Vanky (2002), and for the
new genera consult the mycological literature after 2002.
1h OT Pieridopiyidsi sewer h hg. Dons le ae 2 on ean ot eae Eee eee 2
~ Not on PreridOphy id sas 7 secre aie et ee ae ake ee ee 2
2. On Selaginellaceae; sori as black spots on leaves or stems; spores single, blackish-
brownwembeddedan-the ost tSSue.f ee re er Melaniella
~ On Osmundaceae; sori external on hypertrophied, deformed leaves, cinnamon
brownsispores producediin, chains a. sete oey ee er ae Exoteliospora
3. On Gymnospermae (Araucariaceae); sori in the inflorescence on the surface of
host tissue; one to several spores enclosed by a foveolate coat........... Uleiella
~ On Angiospermae; spores not enclosed by a foveolate coat ................... 4
4, On Eriocaulaceae (seeds) -. 2 ae Bear tts eee ee eee 5
- Notion Brmocailaceqe: ees ot rt ee ar ee eel ee ei gar ene aera 7
5 gr SDOTES SINS eg ek ae. tae a oe ate eRe RN Eriocaulago
- Spotestmsporeiballsae) gyi: et gegen a SN 2 he i cs ee le 6
6. Spore balls mixed with sterile cells and covered by a cortex......... Eriomoeszia
- Spore:balls without. sterile‘cells and.cortext a2 0 Ariapenss Pea See: Eriosporium
7p lwo. ditterent kinds ofispores:presentiin the same sorussseretey eee eae 8
- Spores of the same kind (but conidia or sterile cells may be present)........... 9
8. Small, hyaline, single spores mixed with large, pigmented spore balls; spore mass
grey;'on Cyperacead and juncacedes. wa), wus. ae. eee. oe Heterotolyposporium
D1
2
Ustilaginomycetes — 28 ... 169
Thin-walled, ellipsoidal, ridged spores and thick-walled, globose, tuberculate
spores of the same pale colour present; spore mass butter-coloured; in flowers
OL 120) V PONACE CCH MAr tay ntti ee PA PIAS SIS ia oS ew eh Al Zundeliomyces
Spores mixed with groups of Y-shaped conidia; in ovaries of Poaceae
BAN ee ete ty uc SMM LEO, OREO! acid oy al Dye Conidiosporomyces
Ppores MOLmiIxed. Wit hisuch conidia meenwen mt tanwie lens - 0) Sek. FNM Ee, oem eer, 10
DPOTes Single eee UES oe Lee as 11 (60, 76, 79, 80, 83, 84, 91, 94)
Spores in pairs or in balls, or spores several celled (sometimes in groups or
S01) sa seticas 8 Baird ebewel fie Lge he, Bh senses es wai bce Loris sy sires scans bee RCS 5389)
Spores firmly agglutinated in indefinite masses in the leaves of Convolvulaceae
PY ee see ces Pat piey Bs ee ea ES adaci ess cbse cio, boca tort. & Georgefischeria
Spore wall separated into two layers by a hyaline, gelatinous mass............ 2
Sportewwallinot separatediinto LWormaversmr e973 seta, Le a ee is
Spore wall completely separated into two membranes; the outer membrane
ruptures readily permitting the true spore to pass freely; on Cyperaceae
Ws i Palos pee Te A iulee Ao coe so reely as a nwa MORK Kuntzeomyces
Spore wall separated only in two opposite sides, forming light-coloured polar
Paps sOD CICOts (WallOUs fara ilics mewn an eit ee eae ae oe Yelsemia
Spore mass powdery at maturity, in various parts of the host ................ 14
Spore mass semi-powdery to agglutinated, black; on Cyperaceae and Juncaceae 31
Spore mass not powdery at maturity, spores embedded in the host tissue... ... 40
Sori forming pustular galls on the stems of Tiliaceae .............. Pericladium
DOLLUMOUS OME Aaa PELE Fee ie Me ee! ERI Ee A LR ae, 1
Sori in the flowers of Hyacinthaceae (Liliaceae s.lat.) ............. Antherospora
DOLL SOsme eee see rN nae MN en aC cen me MLL Mere tee oe 16
Spores largeriuaiNlvs( lr) 15-55 Pence ot go eens ee eee ee ee Ae eee, Wy
Sporessinall 10 mediumesized, mainly 5-15 (lens witty vane 3 ee eee Ue
Sori external, forming crusts on leaves of Cyperaceae; spores single or in balls
PM ee RE Sete ee eee eee ae eee Orphanomyces
Sori not external, forming powdery spore masses in the host tissues.......... 18
Sori on leaves forming pustules or swellings; on Liliaceae s.str. ......... Vankya
BOLE Ovabies: NOLO L1iGCEQe! Sen ts as oe Seen Men eet eee ee i
Spore mass golden- to rusty brown; on Cyperaceae and Juncaceae..... Bauerago
Spore mass pale or dark reddish or blackish-brown; on Poaceae ............. 20
Spores elongate, foveolate, with a conspicuous hyaline appendage..... Neovossia
Spores globoid, otherwise sculptured, appendage lacking or short, papilla-like 21
Spores with a dark, pigmented ring; basidia reduced to the spores
170 ...Vanky
22. Spores peculiarly ridged; basidiospores 1-2, ballistosporic....... Ingoldiomyces
- Spores not peculiarly ridged (usually reticulate, verrucose, tuberculate, with
subpyramidal warts or spines, rarely smooth); basidiospores several, not
ballistosporics 2) gaye ot ee bos See oh ee Pee Tilletia
23(16)sSormwithisterile fungalichains or filaments yy eet pase yee eee ee ee 24
- Soripwithout Sterile’chains, Om Dlanientsipe tudo amet eeeee ernie eet mn rns 26
248 HSoriwithisterile tungalichainsonpPogceaen awe ee ee ee Franzpetrakia
- Soriwith filaments;not.onsPogcedenrcean einen: eee ae i es eee ce 25
25. Filaments of uncertain origin; spores violet tinted, produced in cavities in the
flowers or in tumours on the stems of Polygonacede...............00000. Liroa
- Filaments of fungal origin; spores not violet tinted, in flowers of Cyperaceae
2s. spd wap apenbottas Meslks GR age Sree) ait ak) yeti tae seers Farysia
26(23)Omdicots, oc ct eh ck eee ce oe oe See EE Teen ee re re oF
- OnMOonocots.( POGCede) bi. < tate, Gye eee Pee em ater niin ae ee 29
27. On Euphorbiaceae (in seeds of Euphorbia); spore mass brown, without violet tint
set. deo welt or ies ch eraerate 5 etteans ht Core eeoe Ahmadiago —
- Not on Euphorbiaceae; spore mass light to dark purplish-brown, spores violet
tinted o.sxikivstes Wiles Gales greene ath el Meet oe, ieee Te oon ae gee 28
28. Peridium and columella present; spores at first catenate; on Polygonaceae
Bh Date RO: Sates Ski yoo ark: GAG UD eet Veta ane Ra tis, SV oe Sphacelotheca
- Peridium and columella absent; spores not catenate; on various families
SIEGE ROD EE ER CCU rye OO Ie Sr uae BaP Ee Wer eee Sieed Microbotryum
29(26). Sori naked surrounding stems or floral axis; spores usually small, <8 um
ae eT RIE Oe PORTE LE EEL ASS PR ine 6A OR Tranzscheliella
- Sorimot sO: spores usually larger: wie. Seneiece: e Sege ae 30
30. Spores single or in groups of pseudo spore balls, mixed with sterile cells
BELEN SRR yh oer ae, CO AOE RE REE OER, FMP CRN Macalpinomyces
- Spores single, not mixed withustenle cells. see maetsen cn eee ree ee Ustilago
31(13). Sori as onion-like swellings at the base of culms and basal leaves of Eleocharis
Re er See NNT RR AS i ee I PP Ren oo es yeas ae Parvulago
32. Sori around the nuts of Cyperaceae; spores single; basidium 2-celled ......... 33
- Sori in different organs of the host plants in Cyperaceae or Juncaceae; spores
single, rarely in balls, variously sculptured; basidium 4-celled ............... 34
33. Spores with a dark equatorial band and two light polar areas ......... Planetella
- SPOTES NOCSO. cee et eae Ghee teow ete Gt antes Anthracoidea
34. Sori usually with sterile stroma surrounding internodes of the culms or floral
PEdICEls, rarely Ne mpCxOl sclera SpIKCLeLs amie 0. gee meen tee een en meee oa
- DOLL NOMSOie anes Bias yietn GAP is aot eet et etee irc ae tacts ct Ce icc at seine ee 36
Ustilaginomycetes — 28... 171
Sorrpinrection systemic, ost plantsstemlepirmnercy sta. ee lsu: Leucocintractia
- itiecnonnocalised Dostplantsmousterilermes © +6 2 cnt a. eee ee Cintractia
Mam OLUT A SOIC sPikcicts OF alminiOresCellCes, oer r oi an ess te Syne nae eee Oy,
- Sori in all spikelets (or groups of spikelets) of an inflorescence............... 38
37. Sori with a sac-like peridium; spores powdery, mixed with long, hyphal cells
PAE Aer So OITee Rr Ree Ere rE ane Nees thE, EN Sak Trichocintractia
- Sori without sac-like peridium; spores agglutinated, mixed with gelatinised
VDAC res PSIG Men ee eier et Pres cin url PE tee Pilocintractia
Bree ASI LT CACCACsSPOTeS SIN? CON ne Mey ee nes ora ieh oo raienk ee Stegocintractia
- MDG Y DELUCCAC NP Ne te ha OSE ae NGL Oe MER oo. Hic ors Sow ie us ot ae a cod 39
Some TUSCIL DUS, SPOTES SING lente anak bee ruey oe Ana Ls et En eY! Portalia
- On Rhynchospora, Bulbostylis, Cyperus, Scleria; spores single or in balls
PERS ES Dee aie eee Oe: So Es oe ey oe Ustanciosporium
fe) ooritorming irregularcalls pepper weer eee erin tetas at: eerie aoe 4]
~ Sori forming spots, sometimes pustules or rarely swellings or crusts on leaves
ATICESCETIS EES oe eter. Cnet eee renee 9 Roti cae Wc tT eae MME 44
41. Galls multilocular; spores embedded in a gelatinous mass; on Polygonaceae
eet ef lade hale. cere ee erate Lea oh Melanopsichium
- Galls not multilocular, spores not so; not on Polygonacede ...............45. 42
42. Sorias witches’ brooms on leaves or inflorescence of Cyperaceae (Mapanioideae)
with bundles of cylindrical branches; spore mass black; spores not intracellular
Preteens Pieter gn ee atte eh iil ls anal oy oeateks Cintractiella
- Sori as galls on roots (not Mapanioideae); spore mass pale; spores intracellular
nla Ck SS AO OT See MN eae, oR A EC Sey ROE ECP 3PM AER ang ROE A 43
MEMS OULOL (CV Deraccac aud JuncoCcdCe aa a ee fe ee Rie Entorrhiza
- Sori on dicotyledonous plants (in Aizoaceae, Molluginaceae, Portulacaceae)
5 cohen RAUL SAN ee OR ee ane TROT nN entra MeNaN We OR Aone, Talbotiomyces
Bee) spores pigmented, olivaceous or blackish-Drown ao .ee ree 45
- pores pale coloured seebe sia ca geome et Re Pe ae tec 53
45. Sori on dicotyledonous host plants as dark spots, pustules or swellings on leaves,
STems oreLootsi Dasidium Of ul ilictig-(yDC) meme re cn eee ee ee 49
> DOM On mmOnOcOts Om MHOw i Only, (LOU CUI Cyrano eee 46
BOLO RUD DIGCER CA Mera Cte Bd ies ce kaha a) uy si ed yatta aA: Flamingomyces
a POLL MOLONIRUPPIQCEdE, yn. atl ss agentes | oe eiaass Sey cet abicin Aen eee 47
Beem Olt ON POdLede alld OV peraceae NW Ee OR Sines ae ln ee oe 50
= Sori not on Poaceae and Cyperaceae (forming dark leaf spots) ............... 48
48. On Araceae; septal pore simple with two membrane caps and two inner plates
UR tel) bia A ashe Sane Vit oer Pe AE apa MENA TENN ep e Melanustilospora
On Asphodeliaceae, Hyacinthaceae, Melanthiaceae (Liliaceae s.lat.; Eremurus,
Muscari,Narthecium); septal poreaidolipore: aya «euben ote Gjaerumia
172 ...Vanky
49(45). Sori as dark spots on leaves of Fabaceae; septal pore a dolipore, pore caps
Be acters h Sen" wt vedi Dane inet iol weedy, ni ee Wt th Pe ic dt Melanotaenium
50. Basidia 4-celled, phragmobasidia with ballistosporic basidiospores on lateral
SUCEIGTIALAS crete eee easS eM wie len oe ey ond ene ta bin, Ris Seg eT GN Eoue sy GE Tie” aenC sre ee eee Sli
- Basidia 1-celled, holobasidia with apical basidiospores ..................... 52
51. Sori on Poaceae as black leaf spots; spores smooth ........... Phragmotaenium
- Knowdr onlydronr culture; spores ccuinulate «rere ae net ce eee Tilletiaria
52. Sori on Poaceae and Cyperaceae as dark spots, crusts or powdery cover on leaves,
leaf sheaths and stems; ballistospores present ................. Jamesdicksonia
- Sori on Poaceae as dark leaf spots; spores embedded in the leaf tissue; no
ballistOsSpOres = sa x's Rhee sare cae TAN om ner eA eee ween aa Eballistra
53(44). Spore mass powdery, yellowish or pale reddish-brown......... (a few) Tilletia
- Spore mass not powdery, embedded in the host tissue, subhyaline or pale
yellowish- brown f4. 28 tals au wlaceeous to mraieeans 0s cates ore eee 54 |
54. Germination of Ustilago-type; on monocots (Poaceae) .......... Ustilentyloma
_ Germination of [illetia-ty pe; on, dicotsisyiecnet tee se ee eee 55
bd: y eOtraquatic plantst lee asst SPEEA oe eye er re ee ee 56
- Onsterresttial plants: “7 adesnagiehie! see sab aeons eee Entyloma
56. Spores lemon-shaped, with a papilla and an appendage. ........ Rhamphospora
~ Spores globose to broadly ellipsoidal, without papilla and appendage Doassinga
>/(10)opores 11 pairs (rarely the pairs 1McDalls) oc eam a eee eee eee eee 58
~ spores in ballsor'sporesiseveral-celled i was tae ee ene ee 60
58. Spore germination of Mycosyrinx-type; on Vitaceae............... Mycosyrinx
= Spore germination or Usitlavo-ty pe, NOLON Vildccaces s,s ep ae ee aby)
59. Sori hypertrophied; spores produced in cavities of the host tissue; basidiospores
brown:on Sterculiaceaen eas th tt ee Geminago
- Sori not hypertrophied; spores produced intraepidermally; basidiospores
hyaline; om Gyperaccaesyt au via. eins ee Aten, Re ene eee Schizonella
60(57)2 Spores one- to several=celled; on Araliaceaet ae + te Mundkurella
- Spores in ephemeral or permanent balls; not on Araliaceae.................. 61
61. Sori not bursting at maturity, not dusty; spore balls pale, embedded in the host
tissué.ofaquatic.onpaludal plants. 2.72.05 -eer soaker ee ee eee 62
- Sori bursting at maturity, usually granular-dusty; spore balls pigmented ...... 72
62.” "Spore. balls withouvcortextn is oen ns Mae eer cos eee eee, om, 63
- Spore-balisrwith. cortex cies ek no eet tee tele. bem cle a eee oP 65
63. Spore balls consisting of parenchymatous fungal tissue of sterile cells in which
thespores arescatterediy ayes tenets ain ee, eee eS. Burrillia
- No parenchymatous stertle cellsinithe:spore ballspme Aste ene ss vee 2 64
aGki nego rm see at ener se Er a a ee Erratomyces
Sori on various dicots, not on Fabaceae, forming dark leaf spots, pustules or
swellings; septal pore simple with two membrane caps, inner plates lacking
Ustilaginomycetes — 28 ... 173
64. Spore balls loose, friable, consisting of branched hyphae in which the spores are
SU ALLCE CUM em MnennT He Seelyeneee NOPE NE Heer MMe ee SIS LE Se Nannfeldtiomyces
- Spore balls not friable, composed of spores only................. Entylomaster
PMP OrteX Ollitiiny United spotesmemnates ye ete. cess hoe keke Fede e ya ces eee 66
- Sortexopsteriletcell sir nee Renee ee ae tr On ue ore elite earn Une oer, 67
66. Central part of the spore balls consisting of a network of branched hyphae Tracya
~ Central part of the spore balls consisting of parenchymatous fungal tissue
NER es te ih es ie il) tir eee a Me AL, Pseudotracya
Bs spore balls lobed and also containing cavities .....0..-.......5.. Narasimhania
- mpore vallsnoL lobed. compact WiLnOULCaViLieShes as a none en cy Sue... 68
68. Central part of the spore balls composed of hyphae; spores situated between
Lhieseratic: tlie Cortex s Seer shen seed Pe, Mee eae ee PLU ED Pseudodoassansia
- Wenitral partot thespore ballsmot composediofihyphaessse eee. eo. 69
69. Central part of the spore balls composed of parenchymatous sterile cells only;
spores situatedibetweenmtheseand the cortex... eee Doassansiopsis
- Central part of the spore balls composed of spores or also of sterile cells ...... 70
70. Central part of the spore balls composed of spores only -:..-..........-.::- 71
- Central part of the spore balls composed of spores scattered in a network of
STeLile: FUNG AIRCEL Sree ne eters tre ee eee ane A ee ee Pseudodermatosorus
fm Ol tical cells of one kind, notornamented:. ..2..8, 45.20 Fcc ke Doassansia
~ Cortical cells of two kinds: external, small, not ornamented, and internal, larger,
Grnalnentec on stheititinenstiliace per sas eias <0 a ears ee Heterodoassansia
mol spore balls formed by spores and sterile.cells\ ier AM ares. al. eee arse 73
Mee Pore alls rotmed*by spores Only swt viseeee he retina © ott ete settee ach ae 76
meee lhe outer layer of the spore balls is formed by sterile cells’). 7... 0.72.7... ... 74
- The outer layer of the spore balls is not formed by sterile cells ............... v5
74. Spores and sterile cells reticulate; the reticulum between the spores appears as
siaall-empty cells, in seedsiol Cyperacede ete <n e Dermatosorus
- Spores and sterile cells smooth or nearly so; in different organs of host plants of
Warious fanilieses: samen meee en VOT. ET i ey Pet ohne ee eee Urocystis
75. Sterile cells filling the spore balls, surrounded by a cortex of dark spores; on
LED CLOCERE yack aks tint MRR Met het, CPS). OAET oem HERET. ISLE LG UERLYY DR e LAN OF Testicularia
Sterile cells scattered between the spores; cortex lacking; on Poaceae
By fee Pays it ae a ty hice sults tas). ta bles b ete das babes as Moesziomyces
76(72). Between pigmented spore balls and some single spores numerous small,
hyaline, solitary spores present; spore mass grey coloured; on Cyperaceae and
NUP CACE CM wie dete Aare MU Nasties, its oh fu Somervell baer Heterotolyposporium
No hyaline spores between the spore balls; spore mass of another colour...... 77
Spore mass and spores light or dark yellowish, orange- or reddish-brown ..... 78
Spore mass and spores dark, blackish-brown to black ....................5. 84
174 ...Vanky
7825 sterile cells absent between the spore balls yan peer ee eee 79
- Sterile cells usually present between the true or pseudo spore balls; on Poaceae . .82
79. On dicots; spore germination of Thecaphora-type or also with holobasidia
eg A CE RE sate Oa See gens Sa Daiyrs whe. gs 6 Geen me erat ne Thecaphora
~ On monocots; spore germination of Ustilago-type, with phragmobasidia...... 80
80. Spores brown; on Poaceae (spore balls may be ephemeral; sterile cells may be
DLOSCNIL) € rela i iiy Ae ceed ns saint Re ain Aa A Boe cnn Re Sporisorium
- Spores: golden-yellow, on Poncece or Cy peraceac.s., 9.2 ee 81
Sl aaRooruon stems of Poaceae (spore balls persistent) * ate naee eee Fulvisporium
- Sori in spikelets of Cyperaceae (spore balls persistent, loose or spores single)
LS ERS EE MT ek Sar Oe ee TA ADDING HE Fate Roksan nel BED Aurantiosporium
82(78). Two different kinds of sterile cells present between the spore balls; sori
dividedinto compartinentser. ms anare me a arren, ee 2 ees Anomalomyces
~ Only one kind of sterile cells present (or also lacking); sori not divided into
COMP ALIN DUS: 55 hw gare tan get coe este Rees te en ee i erst Ae ee a ee ee 83
83. Spores in true (often ephemeral) spore balls; columella present ....... Sporisorium
- Spores in pseudo spore balls or spores single; columella absent or
tucdimental. *8og veo ne el ere ene, wen ote eae Macalpinomyces
84(77). Fascicles of fungal sterile filaments between the spore balls and spores present;
exospore typically cracked, scaly; on Cyperaceae (Gahnia) ........ Farysporium
- Fascicles of fungal sterile filaments absent; exospore not so.................- 85
Soe OnResiionacede slat ah. ac. We Gee eee ee es a ee eee ee 86
a NOt Of REStONACEQE 2 cia eras ete cia te etn ets aes Sate cee 87
86. Sorias bullate striae on the culms of sterile plants; spores extremely varying in
shape. and 'sizent sera Ae oe te ey ee eer te Websdanea
- SOrl Inthe, seeds sDOles NOLO ae ase et crn cee ee Restiosporium
87. On Centrolepidaceae (seeds of Centrolepis)............... Centrolepidosporium
- Not.on:Centrolepidacedew ee it jrsthe san c wae ea eee Ns ee e e ee 88
88... .sorkin theflowers of*Agavacece (Eniaceae slat). wows. eee ee Floromyces
= SOL NOE SOWA Pate Sat os ns pei ots eels 2d bee de ee 89
89. Sori in the leaf-veins of Rosaceae; spore germination of Ustacystis-type Ustacystis
~ Soriand spore germinationmotso;n0Lom Rosacegesae ne. See eee 90
90. Sori on the surface of young leaves of Cyperaceae (Carex) forming black cover;
spores foveolate or reticulate, in balls or single; basidium short with one, long,
septate basidiospore pea. & She ee a ene seer Orphanomyces
~ SOticsporesiancahasidiamotsoy Ay: sate) ete nike eee ae eee 91
91. Sori on the surface of leaves and inflorescence of Dracaenaceae (Liliaceae s.
lat.; Nolina); spores smooth, in balls or single; basidium bifurcate, with two
basidiospores ttipase: Ae hore ad atl 1 Na, tee Crochet Sateen ots Bartie ieeeee, Clintamra
- Sorispores.and basiddainot $0 gaat paar. kia hak aed Nase oe ey eae 92
Ustilaginomycetes — 28... 175
92. Sori on the surface, or in the spikelets of host plants in the Juncaceae and
Dy PCTACEUERA oP Came tM ORR ee Ne PN, Tolyposporium
- SOU MNOLGU ties See. oe eee is CR. Ce a. es ae Tee ee 8)
93. Sori on the inner, adaxial side of the leaf sheaths of Poaceae; spore wall extremely
varying in thickness; ballisto-basidiospores produced on phragmobasidia
eR ITE Bes Walco, Se Ars soe Sle en OO an aN Tolyposporella
- Sori in the flowers of Cyperaceae; spore wall not so; no ballisto-basidiospores
RLOULICC een en ee aie cesihd, DeerteMtWAS SEE act Ue Re acon ihe (he LE ceuleig 4
94. Spores in firm spore balls; spores deformed, usually wedge-shaped, smooth on
Precontace cides, verrucose On tne freesullace #1. ee ee Moreaua
- Spores in balls or single, usually rounded, not much deformed but slightly
flattened, with two, small, hyaline appendage on the flattened sides, commonly
SRI OY LEAN A a RR ie A UE sak ac Sa al AN LR Ustanciosporium
Acknowledgements
I am grateful to S. Toth (Gédéllé, Hungary) for preparing the Latin diagnoses, to
R.G. Shivas (Brisbane, Australia), and R. Kirschner (Frankfurt, Germany) for reading
the manuscript and serving as pre-submission reviewers, to M. Stoll & D. Begerow
(Tubingen, Germany) for information regarding molecular phylogenetic analyses of
some Tranzscheliella species. The nomenclatural revision is kindly acknowledged for
S. Pennycook (Auckland, New Zealend). Many thanks to L. Guo (Beijing, China) for
the isotype of Doassansiopsis guangdongensis and for unpublished information, to
H. Scholz (Berlin, Germany) for identifying several host plants, to R. Bauer (Tiibingen,
Germany) for unpublished information, to K.-H. Hellmer and M. Wagner-Eha
(Tiibingen, Germany) for technical assistance with SEM pictures, and preparation of
semi-thin slides, respectively. Thanks are also due to the Directors and Curators of the
Herbaria BPI, BR, BRIP, HBG, HCIO, HMAS, IMI and PREM for loans and/or exchange
of specimens.
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Mycol. Balcan. 3:107-118.
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Zundel GL. 1951. Notes on the Ustilaginales of the world. V. Mycologia 43:267-270.
Zundel GL. 1953. The Ustilaginales of the world. Contr. Dept. Bot. School Agric. Pennsylvania State
Coll. 176:XI + 1-410.
MY COTAXON
Volume 106, pp. 179-193 October-December 2008
Psilocybe (Basidiomycotina, Agaricales, Strophariaceae) in Canada,
with a special review of species from British Columbia
GASTON GUZMAN!, PAUL KROEGER?’,
FLORENCIA RAMIREZ-GUILLEN' & RANULFO CASTILLO-DEL-MORAL!’
*gaston.guzman@inecol.edu.mx florencia.ramirez@inecol.edu.mx
Instituto de Ecologia
Apartado Postal 63, Xalapa 91000, Mexico
"kroeger.paul@gmail.com
395 East 40th Avenue, Vancouver, B.C., Canada VS5W 1M1
Abstract — Historically, 28 species of Psilocybe have been reported from Canada, with
the oldest report dating back to 1913. Of these, P corneipes has been transferred to
Mythicomyces corneipes, two (PB. rhombispora, P. subfimetaria) have been synonymized
under earlier names, P callosa has been redetermined as another species, and two
reports (P. merdaria, P. pratensis) are doubtful. With the report of three new records
from British Columbia (PR angustispora, P. apelliculosa, P. subcoprophila), the total
number of verified Psilocybe species recognized in Canada now stands at 24, with the
highest number of taxa concentrated in British Columbia. Eleven of the known taxa of
Psilocybe in Canada are hallucinogenic, with P. quebecensis the only endemic species,
P. semilanceata common throughout the country Canada, and P. pelliculosa apparently
widespread in British Columbia.
Key words — hallucinogenic mushrooms, North America
Introduction
Redhead (1997), who documented macrofungal inventory in British Columbia,
observed, “Our knowledge is most incomplete for macrofungi: less than 1% of
the macrofungi flora of British Columbia have been examined in systematic
studies.” This observation might also reflect what was known about Psilocybe for
most of Canada. MacKay (1913) and Davidson (1930) were the first to report
Psilocybe for the country. MacKay (1913) discussed a case of intoxication in
children in Nova Scotia; the mushrooms in question were sent to Peck, who
identified them as Panaeolus (now Psilocybe) semilanceata (Guzman 1983,
Redhead 1989). Davidson (1930) reported P subviscida from among agarics
collected in Vancouver, BC.
Not until the 1950-—70’s did interest in Canadian Psilocybe species begin to
build, following the discovery of hallucinogenic mushrooms in Mexico, Canada,
and the U.S.A. (Singer & Smith 1958, Olah & Heim 1967, Robinson 1975,
180 ... Guzman & al.
Guzman et al. 1977). The earliest report of recreational use of hallucinogenic
Psilocybe in Canada was in 1965, when the Royal Canadian Mounted Police
confiscated mushrooms — later identified as P semilanceata by A.H. Smith
— from students at the University of British Columbia (Benedict et al. 1967,
Pollock 1978, Davis 1981). Groves (1962) did not include a single Psilocybe
description in his comprehensive Mushrooms of Canada, although he did cite
P. semilanceata “type” species for the genus; Redhead (1979, 1981) subsequently
added a description and an illustration of P semilanceata to the fourth and fifth
editions of Groves’ book. Recently, Nugent & Saville (2004) sequenced DNA
from several species from Canada; included among the species tested was a
culture obtained from a confiscated collection of P cubensis, a tropical species
that is not found in the wild in Canada.
We discuss here all known species of Psilocybe reported in Canada, and
present data from examinations of Psilocybe collections made by the second
author (Kroeger) in British Columbia as well of others from the UBC
Herbarium.
Materials and methods
Bibliographic references on Psilocybe in Canada were extensively reviewed
and Psilocybe collections in the Kroeger personal and UBC herbaria were
studied. Microscopic observations were made of tissues hand sectioned from
basidiomata, mounted in 5% KOH, 1% Congo red and 5% NH,4OH, with at
least 25 measurements made for each microstructure. All drawings were made
with the aid of a light microscope drawing tube. Basidiospore sizes are stated as
length and width in face-view, and thickness in side-view.
Results
From 1913 (MacKay 1913) to the present, 28 species of Psilocybe have been
reported from Canada (TaBLE 1). Verification of these reports required (and
still require) re-examination of herbarium collections because some records
may represent misidentifications. Psilocybe corneipes (Smith 1938, 1949)
was excluded because it belongs to the genus Mythicomyces Redhead & A.H.
Sm. (Redhead & Smith 1986). Collections reported as P rhombispora and P.
subfimetaria belong under the synonymous names P. phyllogena and P. sierrae,
respectively. Collections reported as P callosa have been redetermined as P
strictipes, and P. physaloides probably should be referred to P. subviscida var.
subviscida (Guzman 1995, Noordeloos 2001). The reports of P merdaria
(Lincoff 1981, Stamets 1996) need to be reviewed due to the confusion and
the lack of herbarium material; they probably also represent another species.
Three species are reported for the first time from Canada, as discussed below
(see TABLE 1).
Psilocybe in British Columbia (Canada) ... 181
We now recognize 24 species of Psilocybe for Canada, which represents 11% of the
227 species reported worldwide by Guzman (2005). There are 11 hallucinogenic
species found in Canada, which represent 8% of the 144 accepted by Guzman
(2005) in the world. Most Psilocybe species known from Canada occur in British
Columbia, with only a few reported from eastern Canada, mainly in Ontario
(TABLE 1). Of the hallucinogenic species in Canada, P. quebecensis appears
to be the only endemic and P. semilanceata the most widespread in Canada;
P. pelliculosa is commonly reported in British Columbia.
TABLE 1. Known species and new records of Psilocybe in Canada '
*P. acadiensis A.H. Sm., J. Elisha Mitchell Sci. Soc. 62: 192, 1946. NS (Smith & Hesler 1946, Guzman
1983).
*P. angustispora A.H. Sm., J. Elisha Mitchell Sci. Soc. 62: 193, 1946. BC.
*P. apelliculosa P.D. Orton, Notes R. Bot. Gard. Edinb. 29: 118, 1969. BC.
*P. atrobrunnea (Lasch) Gillet, Les Hyménomycetes, p. 586, 1878 (Redhead 1979, 1981, Stamets
1996; without locality).
P. baeocystis Singer & A.H. Sm., Mycologia 50: 141, 1958. BC (Robinson 1975, Guzman et al. 1977,
1998, Repke et al. 1977, Stamets 1978, Guzman 1983, Ammirati et al. 1985, Redhead 1997).
*P. bulbosa (Peck) A.H. Sm., Mycologia 40: 694, 1948. ON (Redhead 1984).
P. caerulipes (Peck) Sacc., Syll. Fung. 5: 1051, 1887. ON (Singer & Smith 1958, Robinson 1975,
Ammirati et al. 1985, Guzman et al. 1998).
[P. callosa (Fr.) Quél., sensu Guzman 1983 (see P. strictipes) |
*P. coprophila (Bull.) P. Kumm., Fuhr. Pilzk., p. 71, 1871, BC, QC (Guzman 1983).
[*P. corneipes (Fr.) P. Karst., sensu A.H. Smith (1938, 1948, 1949) = Mythicomyces corneipes (Fr.)
Redhead & A.H. Sm., Can. J. Bot. 64: 643, 1986. BC, NS, ON (Smith 1938, 1948, 1949, Stamets
1978, Redhead & Smith 1986, Huhtinen & Vauras 1992)].
P. cyanescens Wakef., Trans. British Myc. Soc. 29: 141, 1946. BC (Robinson 1975, Guzman 1978,
1983; Stamets 1978, 1996, Lincoff 1981, Ammirati et al. 1985, Redhead 1997, Guzman et al.
1998).
P. cyanofibrillosa Guzman & Stamets, Mycotaxon 11: 477, 1980. BC (Stamets 1996, Guzman et al.
1998).
P. fimetaria (P.D. Orton) Watling, Lloydia 30: 150, 1967. BC, NB (Guzman 1983, Stamets 1996,
Redhead 1997, Guzman et al. 1998).
*P. inquilina (Fr.) Bres., Icon. Mycol. 18, pl. 863, 1931. BC (Kroeger 1989, Redhead 1997).
[*P. merdaria (Fr.) Ricken, Blatterpilze, p. 251, 1912. BC, QC (Lincoff 1981, Guzman 1983, Stamets
1996)].
*P. montana (Pers.) P. Kumm., Fuhr. Pilzk, p. 71, 1871, without locality (Redhead 1979, 1981).
P._ pelliculosa (A.H. Sm.) Singer & A.H. Sm., Mycologia 50: 280, 1958. BC (Robinson 1975, Guzman
et al. 1977, 1998, Repke et al. 1977, Redhead 1979, 1981, 1997, Lincoff 1981, Guzman 1983,
Ammirati et al. 1985).
' {information in brackets] = excluded or questionable records; * = non-hallucinogenic species; bold-face =
new records
Canadian provinces (abbreviations): Alberta (AB); British Columbia (BC); Manitoba (MB); New Brunswick
(NB); Newfoundland (NL); Nova Scotia (NS); Ontario (ON); Prince Edward Island (PE); Quebec (QC);
Saskatchewan (SK).
182 ... Guzman & al.
TABLE 1, concluded
*P. phyllogena (Peck) Peck, Bull. N.Y. State Mus. 157: 99, 1912. BC, NB (Guzman 1983, Redhead
1997, as P. rhombispora).
[*P. physaloides (Bull.) Quél., Mem. Soc. Emul. Montbéliard (Ser. 2) 5: 256, 1872, nomen dubium;
see discussion of P. subviscida var. subviscida. |
[*P. aff. pratensis P.D. Orton, Notes R. Bot. Gard. Edinb. 29: 120, 1969. BC (Kroeger 1989, Redhead
1997)].
P. quebecensis Olah & R. Heim, Compt. Rend. Hebd. Séances Acad. Sci. 264: 1601, 1967. QC
(Olah 1967, 1973, Olah & Heim 1967, Guzman 1978, 1983; Stamets 1978, 1996, Chilton 1978,
Guzman et al. 1998).
[*P. rhombispora (Britzelm.) Sacc., Syll. Fung. 11: 72, 1895, a synonym of P phyllogena}.
*P. sabulosa Peck, Bull. Torrey Bot. Club 24: 144, 1897. BC (Redhead 1997).
P. semilanceata (Fr.) P. Kumm., Fuhr. Pilzk., p. 71, 1871. BC, NB, NL, NS, PE, QC (Groves 1962,
Heim et al. 1966?, Olah 1967, Robinson 1975, Guzman et al. 1977, 1998; Repke et al. 1977, Weil
1977, Stamets 1978, 1996, Lincoff 1981, Guzman 1983, Ammirati et al. 1985, Redhead 1979,
1981, 1989, 1997, Schalkwijk-Barendsen 1991a, b, Turner & Szczawinski 1991).
*P. sierrae Singer, Beih. Nova Hedwigia 29: 240, 1969. BC (Guzman & Smith 1978, Guzman 1983,
1995, Stamets 1996, Redhead 1997, Guzman et al. 1998; some as P. subfimetaria).
P. silvatica (Peck) Singer & A.H. Sm., Mycologia 50: 277, 1958. ON (Singer & Smith 1958, Robinson
1975, Stamets 1978, 1996, Redhead 1979, 1981, Guzman et al. 1998).
P. strictipes Singer & A.H. Sm., Mycologia 50: 141, 1958 [= P. callosa (Fr.) Quél., sensu Guzman
1983]. BC (Robinson 1975, Guzman et al. 1977, 1998, Guzman 1983, 1995, Ammirati et al.
1985, Stamets 1996, Redhead 1997).
P. stuntzii Guzman & J. Ott, Mycologia 68: 1261, 1977. BC (Guzman & Ott, 1977, Guzman et
al. 1977, 1998, Repke et al. 1977, Stamets 1978, 1986, Guzman 1983, Ammirati et al. 1985,
Redhead 1997).
*P. subcoprophila (Britzelm.) Sacc., Syll. Fung. 11: 72, 1895. BC.
[*P. subfimetaria Guzman & A.H. Sm., Mycotaxon 7: 518, 1978, a synonym of P. sierrae)|
*P. subviscida (Peck) Kauffman var. subviscida, Agar. Mich., p. 275, 1918. BC, NB (Davidson 1930,
Lowe 1969; Guzman 1983, Stamets 1996, Redhead 1997; some as P. physaloides).
Discussion of British Columbian Psilocybe collections
Psilocybe angustispora (Fics. 1-2)
SPECIMEN STUDIED. BRITISH COLUMBIA, Saturna Island, on deer pellets, September
18, 1983, Kroeger 590 (UBC-2730).
BASIDIOSPORES 11-14 x 6-7 x 6-7 um, subellipsoid, thick-walled, wall 1-1.5
um thick, with a broad germ pore; PLEUROCYSTIDIA absent; CHEILOCYSTIDIA
30-42 x (6-)7-10 um, lageniform.
ComMENnts: Psilocybe angustispora is close to P. subcoprophila, but the latter has
longer spores — (13-)14-18(-—22) um (Guzman 1983).
This species was known only from the U.S.A. (Colorado, Idaho, Oregon and
Washington) (Smith & Hesler 1946, Guzman 1983, Stamets 1996). It is defined
by its mycenoid basidiomata and its peculiar habitat on dung, mainly pellets
of deer and elk, but also grows on horse dung, in temperate forests or alpine
meadows. ‘This report is the first record for Canada.
Psilocybe in British Columbia (Canada) ... 183
Psilocybe apelliculosa (Fics. 3-6)
SPECIMENS STUDIED. BRITISH COLUMBIA, Vancouver, Sept. 22, 1981, Kroeger 01;
Botanical Gardens of the University of British Columbia, Asian Gardens, September
12, 1985, Kroeger 868, 869; September 19, 1987, Kroeger 1251. University of British
Columbia campus, September 18, 1997, Kroeger 2063; September 21, 1997, 2064, 2065
(all in Kroeger Herbarium, UBC and XAL; 01, 868, 869 & 1251 also in DAOM).
PILEUS (6—)10-20 mm diam, convex to subumbonate, finally plane, slightly
viscid to dry, hygrophanous, with silky and grayish fibrils, dark cinnamon-
brown or dark reddish-brown to pale reddish-brown or pale orangish on the
margin. LAMELLAE adnate to short decurrent, grayish when young to brownish-
gray or dark brown-violaceous when mature, edges whitish and subfloccose or
denticulate, finally concolorous or slightly paler. Stripe (17—)30-40 x 1.5-2.5
mm, equal or slightly thickened at apex, hollow, often flexuous, base subfloccose,
often extending into the substrate and covered with white tomentum; surface
dark gray-brown to ochraceous-honey, finally reddish-brown, sometimes
covered with floccose white scales. VEIL cortinaceous, fugacious, remains as
adpressed pale yellowish fibrils in the upper part of the stipe and in the pileus
margin. BASIDIOSPORES (5—)5.5-7(-8) x 4-5 x 3.5-4.5 um, subrhomboid or
subellipsoid in face-view, subellipsoid in side-view, thin-walled, wall up to 0.5
um thick, yellowish-brown. PLEUROCYSTIDIA absent. CHEILOCYSTIDIA (13.5-
)18-31 x (6-)8-12(-14) um, utriform, broadly utriform or sublageniform,
hyaline. PILEIPELLIs with prostrated hyaline hyphae, not gelatinous.
HasirTatT: Scattered to gregarious or caespitose, in dense troops on soils
supplemented with wood-chips-manure compost, in recently establish
horticultural plantations and lawns.
ComMENTs: The above description agrees with those provided by Orton (1969)
and Guzman (1983). Noordeloos (2001) regards P. apelliculosa as a synonym
of P. castanella; Guzman (1983), however, notes that these species differ in the
width of the cheilocystidia [3-5(-6) um in P. castanella].
Psilocybe apelliculosa is a European taxon previously unknown from America
(Guzman 1983). This report is the first for Canada and North America, where
its occurrence in recently cultivated areas (i.e., the University of BC Botanical
Gardens) suggests it has been imported European or Asiatic plants. Psilocybe
apelliculosa is distinguished by its pileus without a separable pellicle and its
wide cheilocystidia.
Psilocybe baeocystis (Figs. 10-11)
SPECIMENS STUDIED. BRITISH COLUMBIA, Vancouver, Downtown, November 28,
1981, Kroeger 89 (UBC-10235).
BASIDIOSPORES (9—)10—13(-15) x (5-)5.5-6.5(-7) um, thick-walled, wall up to
1 um thick; PLEUROCYSTIDIA absent; CHEILOCYSTIDIA 19-33(-40) x (5-)6-8
um, sublageniform, frequently and irregularly branched.
184 ... Guzman & al.
ON
PiaTE I. Fics. 1-10. Canadian species of Psilocybe. 1-2: PB angustispora, 1 basidiospores,
2, cheilocystidia (both from Kroeger 590), 3-6: P. apelliculosa, 3 basidiomata (Kroeger 2064),
4 basidiospores, 5-6 cheilocystidia (4-5 Kroeger 868; 6 Kroeger 869), 7-9: P. coprophila,
7-8 basidiospores, 9 cheilocystidia (7 Kroeger 2188; 8-9 Kroeger 2198). 10: P baeocystis,
basidiospores (Kroeger 89). Scale bar 1-2 & 4-10: 10 um, 3: 15 mm.
CoMMENTs: Psilocybe baeocystis, a common hallucinogenic fungus in British
Columbia (see TaBLE 1), was originally described from Oregon (U.S.A.)
by Singer & Smith (1958) where it is also very common (Guzman 1983).
This species is distinguished by its bluing robust basidioma and strongly
hygrophanous pileus, as well as its asymmetrical subellipsoid basidiospores,
and growth on coniferous bark mulch in gardens and parks.
Psilocybe in British Columbia (Canada) ... 185
Psilocybe coprophila and P. merdaria (Fras. 7-9)
SPECIMENS STUDIED. BRITISH COLUMBIA, Vancouver, April 15, 1998, Kroeger 2184
(UBC-14468); May 10, 1998, Kroeger 2188 (UBC-14542); May 17, 1998, Kroeger 2198
(UBC-14500).
ANNULUS & VEIL absent; BASIDIOSPORES (9—) 10-13 tm long, which agree well
with P. coprophila.
ComMENTs: Both P. coprophila and P. merdaria are fimicolous mushrooms with
subhexagonal, thick-walled basidiospores. Psilocybe merdaria is distinguished
by its well-formed ring on the stipe and the appendiculate pileus margin
and P coprophila is distinguished by the lack of ring and poorly developed
veil (Guzman 1983). These fungi are very common everywhere, although
P. merdaria is more frequent in temperate regions and P. coprophila is typically
tropical or subtropical in distribution (Guzman 1983). The records of P. merdaria
and P coprophila from Canada are rare and contradictory. Guzman (1983)
mistakenly based a report of P merdaria and P. coprophila from Quebec on the
same collection (Cooke 31597, MICH), which is in need of re-examination.
Two other P merdaria reports by Lincoff (1981) and Stamets (1996) also need
verification.
Psilocybe cyanescens (Figs. 12-14)
SPECIMENS STUDIED. BRITISH COLUMBIA, Vancouver, urban zone, November 2,
1976, Guzman 16630 (ENCB). University of British Columbia, Endowment Lands,
October 11, 1981, Kroeger 156 (UBC-10241).
BASIDIOSPORES 11-12(-16) x (6—)6.5-7(-8) x (6-)6.5-7 um, ellipsoid, thick-
walled, wall up to 1.5 um thick; PLEUROCYSTIDIA 17-30 x (4-)5.5-8(-9) um,
common, clavate or narrowly subglobose, both with a short cylindrical apex;
CHEILOCYSTIDIA 18-29 x 5.5-7 um, sublageniform or narrowly utriform,
sometimes branched.
ComMENTSs: This species, like P baeocystis, is another common bluing and
hallucinogenic mushroom from British Columbia (see TABLE 1). P cyanescens,
originally described from England, appears to represent a complex. In fact the
presence of pleurocystidia in the North American taxon suggests it may be a
separate species, as in Europe P cyanescens has no pleurocystidia (Watling &
Gregory 1987, Guzman 1983, Krieglsteiner 1986). The senior author is now
revising this complex.
Psilocybe cyanofibrillosa (Frcs. 15-20)
SPECIMENS STUDIED. BRITISH COLUMBIA, Meager Mountains, Upper Lillooet River
Gorge below Plinth Peak, July 22, 1989, Kroeger 1401; August 16, 1989, Kroeger 1403,
1404, 1405; September 17, 1995, Kroeger 1790; September 1351995, Kroeger 1791, 1792;
1793 [all in Kroeger Herbarium and UBC, except 1404 which is also in XAL and DAOM;
1401, 1403 and 1405 also in DAOM J.
186 ... Guzman & al.
Pate II. Fics. 11-17. Canadian species of Psilocybe. 11: P. baeocystis, cheilocystidia (Kroeger 89).
12-14: P. cyanescens, 12 basidiospores, 13 pleurocystidia, 14 cheilocystidia (all from Kroeger 156).
15-17: PB. cyanofibrillosa, 15 basidiomata (Kroeger 1403), 16 basidiospores, 17 cheilocystidia (both
from Kroeger 1401). Scale bar 11-14 & 16-17: 10 um, 15: 10 mm.
Prteus (5—)10-25 mm diam., conic to conic-convex or campanulate, viscid,
with a well-developed gelatinous pellicle, reddish-brown to pale ochre,
sometimes with a bluing margin. LAMELLAE sinuate, with a decurrent tooth,
pale ochraceous to violaceous-ochraceous, with white edges. StripE 20-55 x
1.5-3 mm, equal, whitish to pale ochraceous, bluing at the base, smooth above,
but covered by white to grayish adpressed silky little scales from the veil to
the base, hollow to stuffed with whitish mycelium. VEIL white and arachnoid
in young stages, the remains forming small white scales on the pileus margin
Psilocybe in British Columbia (Canada) ... 187
and white fibrils on the upper part of the stipe, forming a scaly or floccose
ring. CONTEXT pale ochraceous, bluing. BAsip1osporEs (9-)10-12(-16) x 5-
6(-6.5) x 5-6 um, subellipsoid both in face- and side-view, thick-walled, wall
up to 1 um thick. PLEUROcysTIpIA absent or sometimes scarce, 14.5-16 x 6-8
lum, ventricose, subglobose-mucronate or subglobose-rostrate to subfusiform
or sublageniform, hyaline. CHEILOCysTIDIA (16-)18-32 x (4—-)5-7(-8) um,
sublageniform, frequently irregularly branched with two long necks, hyaline,
sometimes with a dark oily drop at the apex. SUBHYMENIUM with hyaline to
yellowish hyphae, 2-4 um wide. PILEIPELLIS as an ixocutis, 70-120 um thick,
with hyaline hyphae, 1.5-4 um wide. SuBPELLIs with inflated hyaline hyphae.
CLAMP CONNECTIONS present.
HaBiTaT: Scattered to gregarious, in mossy sandy soil on tangled roots of
Alnus, Populus and Salix, frequently inundated flat areas beside river, never in
meadows.
COMMENTs: Stamets et al. (1980) first described this bluing species from
Washington state, U.S.A. and later (Stamets 1996) reported it from northern
California to British Columbia. Psilocybe cyanofibrillosa appears common in
British Columbia, mainly in Lillooet River region. It is close to PR. cyanescens
and is placed in section Semilanceatae based on its subellipsoid, thick-walled
spores (Guzman 1995). A well-developed veil that produces a scaly or floccose
ring and the lack or rarity of pleurocystidia help separate P cyanofibrillosa
from P. cyanescens (Stamets 1996, Guzman 1995). P. cyanofibrillosa differs
from P. semilanceata in shorter basidiospores [(11—)12-15(-18) um long in P.
semilanceata] and the presence of veil (poorly developed in P. semilanceata,
Guzman 1983).
Psilocybe montana (Figs. 21-22)
SPECIMEN STUDIED. BRITISH COLUMBIA, Pasayten Valley, October 17, 1981, Kroeger
164 (UBC-10255).
BASIDIOSPORES 7-8.5(-9) x (4—)5-5.5 x 4-5 um, subrhomboid or subellipsoid
in face-view, subellipsoid in side-view, thick-walled, wall up to 1 um thick,
dark yellowish-brown, with a broad germ pore; CHEILOCYSTIDIA 17-26 x (4-
)6-7 tum; PLEUROCYSTIDIA absent. PILEIPELLIS an ixocutis up to 10 um thick,
composed of 1.5-3 um wide, hyaline hyphae.
ComMENTs: Psilocybe montana, a non-bluing fungus occurring on mosses, is
widespread in temperate regions (Guzman 1983, 1995; Noordeloos 2001) but
rarely reported from Canada, although Redhead (1979, 1981) cited this species
indirectly. The studied BC material agrees with the classic concept of the species
except for smaller cheilocystidia, cf. (15-)22-45 x 4-7.5(-10) um (Guzman
1983) or 17-46 x 4-8.5 um (Noordeloos 2001). More collections should be
examined in order to clarify the cheilocystidial variation.
Psilocybe in British Columbia (Canada) ... 189
Psilocybe pelliculosa (Fics. 23-24)
SPECIMENS STUDIED. VANCOUVER, Shaughnessy, November 15, 1980, Kroeger 51
(UBC-10231).
BASIDIOSPORES 10-13 x 6-7 um, ellipsoid, thick-walled, wall up to 0.8 um;
PLEUROCYSTIDIA absent; CHEILOCYSTIDIA 19-37 x 4.5-6.5 um, lageniform,
frequently irregularly branched.
CoMMENTs: This bluing hallucinogenic species is common in British Columbia
(see TABLE 1). Psilocybe pelliculosa resembles P. semilanceata, for which it is
frequently mistaken — both species stain slightly blue. Smaller basidiospores
and pileus that is neither conic nor papillate distinguishes P pelliculosa.
Psilocybe stuntzii (Fics. 25-26)
SPECIMEN STUDIED. VANCOUVER, University of British Columbia campus in
Endowment Lands, in woodchips, October 25, 1979, Kroeger 42 (UBC-10242).
BASIDIOSPORES 9.5-11 x 6-7 um, subellipsoid or subrhomboid in face-view,
subellipsoid in side-view, thick-walled, wall up to 1 um thick; PLEUROCYSTIDIA
absent; CHEILOCYSTIDIA 20-32 x 4-6.5(-8) um, sublageniform, frequently
irregularly branched.
ComMENTs: A common bluing and hallucinogenic collybioid mushroom
found in urban or suburban habitats, P stuntzii is diagnosed by its annulus and
subrhomboid basidiospores. The collection above agrees well with Guzman &
Ott (1977) and Guzman (1983).
Psilocybe subcoprophila (Fics. 27-28)
SPECIMEN STUDIED. BRITISH COLUMBIA, Saturna Island, April 22, 1984, Kroeger
185 (UBC 12215).
BASIDIOSPORES 14-18 um long, subellipsoid and thick-walled; CHEILOCYSTIDIA
30-45 x 6-10 um, sublageniform; PLEUROCYSTIDIA absent.
COMMENTS: As discussed above, this species is close to P angustispora, from
which differs in its bigger basidiospores. It is a fimicolous European species, until
recently known elsewhere only from Argentina and Greenland (Guzman 1983).
Guzman & Trappe (2005) reported it from an alpine habitat in Washington,
U.S.A. The collection above was found on bovine dung in a meadow.
PrateE III. Fics. 18-30. Canadian species of Psilocybe. 18-20: P. cyanofibrillosa, 18 basidiospores,
19 pleurocystidia, 20 cheilocystidia (all from Kroeger 1790). 21-22: P. montana, 21 basidiospores,
22 cheilocystidia (both from Kroeger 164). 23-24: P. pelliculosa, 23 basidiospores, 24 cheilocystidia
(both from Kroeger 51). 25-26: P. stuntzii, 25 basidiospores, 26 cheilocystidia (both from Kroeger
42). 27-28: P. subcoprophila, 27 basidiospores, 28 cheilocystidia (both from Kroeger 185).
29-30. P. subviscida var. subviscida, 29 basidiospores, 30 cheilocystidia (both from Kroeger 1659).
Scale bar 10 um.
190 ... Guzman & al.
Psilocybe subviscida var. subviscida vs. P. physaloides (Fics. 29-30)
SPECIMEN STUDIED. BRITISH COLUMBIA, Vancouver Island, Lake Cowichan
Research Forest, Mesachie Lake, Mesachie Hill, May 9, 1993, Kroeger 1659. (Kroeger
Herbarium, UBC and XAL).
BASIDIOMA collybioid; PILEUS 6-20(-25) mm diam, convex to campanulate
or subplane, smooth, hygrophanous, reddish-brown to pale yellowish-brown
(straw color), subviscid to dry, without a peelling pellicle, margin translucent
striate when moist. LAMELLAE subadnexed or sinuate, reddish-brown, with
concolorous edges, which are serrate or fimbriate. SrrpE 20-50 x (=) Sena
cylindrical, subbulbous, flexuous, hollow, reddish-brown as the pileus, blackish
at the base, covered thinly with brownish-grey fibrils. Veil poorly developed.
CONTEXT whitish, but brownish-red below the pileus and stipe surface. White
cordons of mycelium are common at the base.
BASIDIOSPORES 6.5-7 x 4.5—-5 x 4-5 um, subrhomboid or subovoid in face-
view, subellipsoid in side-view, thin-walled, wall up to 0.5 um thick, yellowish-
brown, with a broad germ pore. PLEUROCySTIDIA absent. CHEILOCYSTIDIA
22.5-34.5 x 5-7 um, sublageniform. PILEIPELLIS subgelatinous, 10-20 um —
thick, with hyaline hyphae, 1-3 um wide.
COMMENTS: Smith (1948) and Noordeloos (2001) both examined the holotype
of P. subviscida cite different measurements for the basidiospores — 7-8 x 4-
4.5(-5) um by Smith, (6-)6.5-7.5(-8) x 4.5-5(-5.5) um by Noordeloos — and
cheilocystidia — 30-40 x 6-7 ttm or 20-26 x 10-12 um by Smith, 27-44 x 6-8
uum or 23-45 x 3.5-10 um by Noordeloos.
Psilocybe subviscida is a common North American and European
species (Guzman 1983, Noordeloos 2001) that is sometimes confused with
P. physaloides. This latter species has different interpretations by Fries, Ricken,
Bresadola, Cooke, J. Lange, Orton, and Watling & Gregory (Noordeloos 2001),
and Guzman’s (1983) concept of P. physaloides appears to represent P subviscida.
With Noordeloos (2001), we agree that P physaloides is a nomen dubium and
now refer previous New Brunswick (Canada) records of P. physaloides (Guzman
1983, Stamets 1996) to P. subviscida (TABLE 1). Davidson (1930), Lowe (1969)
and Redhead (1997) also have reported P. subviscida from British Columbia.
Acknowledgments
The authors thank Laura Guzman-Davalos, Universidad de Guadalajara, and Timothy
J. Baroni, State University of New York, College at Cortland, for kindly reviewing this
paper. Guzman, Ramirez-Guillén and Castillo-Del Moral thank also Etelvina Gandara,
Juan Lara Carmona and Manuel Hernandez, from Instituto de Ecologia at Xalapa, for
their assistance in laboratory, herbarium and computational duties, respectively. The
senior author expresses thanks to Instituto de Ecologia, CONACYT and SNI for their
support of his research. Kroeger expresses thanks to Scott Redhead for his assistance, to
Psilocybe in British Columbia (Canada) ... 191
the Botany Department of the University of British Columbia for access to facilities and
resources, the Campbell family for hospitality on Saturna Island, and to Trudy Greif for
assistance. The Herbarium of the University of British Columbia is acknowledged for
providing a loan of its Psilocybe collections.
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Stamets P. 1978. Psilocybe mushrooms and their allies. Homestead Book, Seattle.
Stamets P. 1996. Psilocybin mushrooms of the world. Ten Speed Press, Berkeley.
Stamets P, Beug MW, Bigwood JE, Guzman G. 1980. A new species and a new variety of Psilocybe
from North America. Mycotaxon 11: 476-484.
Turner NJ, Szczawinski AF. 1991. Common poisonous plants and mushrooms of North America.
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Watling R, Gregory NM. 1987. 5/Strophariaceae & Coprinaceae p.p. In British Fungus Flora, Agarics
and Boleti. Royal Botanic Garden, Edinburgh.
Weil AT. 1977. The use of psychoactive mushrooms in the Pacific Northwest: an ethnopharmalogic
report. Botanical Museum Leaflets of Harvard University 25: 131-149.
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MYCOTAXON
Volume 106, pp. 195-198 October-December 2008
Diplomitoporus allantosporus (Basidiomycetes):
a new record for Brazil
ALCINDO MARTINS-JUNIOR’, TATIANA GIBERTONI*® & HELEN SOTAO’
micologo13@hotmail.com, tbgibertoni@hotmail.com, helen@museu-goeldi. br
* Museu Paraense Emilio Goeldi, Coordenacdo de Botanica
Caixa Postal 399, CEP 66040170, Belém, PA, Brazil
? Universidade Federal de Pernambuco, Departamento de Micologia
Av. Nelson Chaves s/n, CEP 50760-420, Recife, PE, Brazil
Abstract — Diplomitoporus allantosporus, collected in the Ferreira Penna Scientific
Station in the state of Para of Brazilian Amazonia, is reported as a new record to the
country and the second of the species. A description of D. allantosporus and a key to the
accepted species of the genus in Brazil are provided.
Key words — Polyporaceae, taxonomy, diversity
Introduction
Diplomitoporus is a cosmopolitan genus described by Domanski (1970)
characterised by its resupinate to effused-reflexed, light-coloured basidiomata,
di-trimitic hyphal system, clamped generative hyphae, thick-walled, non-
amyloid to weakly amyloid skeletal hyphae, and IKI- allantoid to ellipsoid
basidiospores (Ryvarden & Gilbertson 1993). Approximately 16 species are
currently recognized for the genus, 14 of which occur in the Americas (www.
indexfungorum.org, www.cbs.knaw.nl, Coelho 2008a, b).
Recently four species have been recognized as new to Brazil: Diplomitoporus
dilutabilis (Loguercio-Leite & Wright 1998), D. taquarae (Coelho 2008a), and
| D. marianoi-rochae (Coelho 2008b) from Southern Brazil and D. navisporus
(Gibertoni et al. 2004) from Northern Brazil. Until now, these were also the
only Diplomitoporus species known for the country. The present paper adds
Diplomitoporus allantosporus, a recently described species known only from
_ its type locality in Venezuela (Ryvarden & Iturriaga 2003). A description of D.
| allantosporus based on Brazilian material accompanied by a key to Brazilian
Diplomitoporus species is presented below.
196 ... Martins-Junior, Gibertoni & Sotao
Material and methods
The Ferreira Penna Scientific Station (ECFPn) is a 33.000ha reserve managed
by the Museu Paraense Emilio Goeldi (MPEG) and is located in the Amazonian
State of Para, 400km from Belém, Parda’s capital city. This reserve is composed
of typical Amazonian ecosystems with a flora that is among the richest and
densest in the Amazonian basin (Lisboa 1997, 2002). The many fungi that have
been collected there during the last 10 years are part of the large collection now
deposited at MG (Sotao et al. 1997, 2002).
The basidioma was analysed macro- (i.e., shape, colour, hymenial surface)
and micro-morphologically (i.e., hyphal system, presence/absence and measures
of sterile structures and basidiospores). Microscopical observations were made
from slide preparations mounted in 5% KOH and stained with 1% aqueous
phloxine and/or Melzer’s reagent (Ryvarden 1991). Colour designation follows
Watling (1969).
Taxonomy
Diplomitoporus allantosporus Ryvarden & Iturr., Mycologia 95(6): 1067, 2003.
BASIDIOMATA annual, resupinate. HYMENIAL SURFACE Saffron (49) to luteous
(51); pores angular, 2-3/mm. Context thin.
HyYPHAL SYSTEM dimitic, generative hyphae hyaline, 3-5 um diam., clamped,
thin to slightly thick-walled; skeletal hyphae hyaline to yellow, 4-5 um in diam.,
thick walled. Basrpia clavate, 10-12 x 3-5 um. Basrprospores allantoid,
smooth, 5-6.25 x 1.5-2.5 um, thin-walled, occasionally guttulate, hyaline,
IKI-.
MATERIAL EXAMINED: BRAZIL. State of Para: Municipality of Melgaco, Floresta Nacional
de Caxiuana, ECFPn (51’27"17° W, 144°12° S), 23.1X.1996, H. Sotdo et al. 96-41 (MG).
REMARKS: Diplomitoporus allantosporus is diagnosed by the size and shape
of its basidiospores. It resembles D. venezuelicus Ryvarden & Iturr., which is
differentiated by smaller (4-4.5 x 1.2-1.5 um), cylindric basidiospores and
smaller, round pores, 6-8/mm (Ryvarden & Iturriaga 2003). The Brazilian
species D. dilutabilis has angular to elongated pores, trimitic hyphal system
and cylindrical to slightly allantoid basidiospores (Loguercio-Leite & Wright
1998), while D. navisporus has also round, regular pores, but trimitic hyphal
system and navicular basidiospores (Gibertoni et al. 2004). Diplomitoporus
taquarae is distinguished by smaller pores [(2—) 4-5 (-6)/mm] and narrower
generative hyphae [(1.8-) 2-2.8 (-3.2) um diam.] (Coelho 2008a), while
D. marianoi-rochae has larger pores [(0.5-) 1-2/mm], branched contextual
skeletal hyphae, broader spores (3.5-9.5 x 1.5-3 um), and abundant hyphidia
or dendrohyphidia when fresh (Coelho 2008b). This represents the second
occurrence of D. allantosporus, previously known only from its type locality in
Venezuela, and the first report for Brazil.
Diplomitoporus allantosporus, new to Brazil ... 197
Working key to Brazilian species of Diplomitosporus
BeeVORALS Vsti tLiTMitiC apie et sm eeens Pe cage tn tek Sgt wae Pan aa ve seams 2
Beem Dia cy sternite aan Mere Meee OPN aes WS ccc so Me ees creel ah dees ee 3
2a. Basidioma resupinate to effused-reflexed, pores 4—6/mm, angular to radially
elongated, basidiospores cylindrical to slightly allantoid, 4.5-5.5 x 2.5-3 um
pei 215 5 OR A UPS oor righ dn, Tera 5 CR OP OPE EP eee MES EEE D. dilutabilis
2b. Basidioma resupinate, pores 6-7/mm, round, basidiospores cylindrical to
AVQWACU LAT Ao) seed Da) een ere ee eee ue tn opts ote Sonne iete D. navisporus
3a. Pores up to 2/mm, hyphidia or dendrohyphidia present ...... D. marianoi-rochae
3b. Pores smaller, more than 2/mm, hyphidia or dendrohyphidia absent ........... 4
4a. Basidioma resupinate, pores 2-3/mm, generative hyphae 3-5 um diam. with
large clamps, basidiospores 5—6 x1.2-1:5)imi). 33.4... D. allantosporus
4b. Basidioma resupinate, soon reflexed at the margins, pores 4—5/mm, generative
hyphae 2-2.8 diam., basidiospores 4.4-6.4 x 1.2-1.8um .......... D. taquarae
Acknowledgments
The authors would like to thank Mario Rajchenberg and Annarosa Bernicchia for
critically reviewing the manuscript; the staff of the ECFPn and of the MPEG for support;
Conselho Nacional de Desenvolvimento Cientifico (CNPq) for the master scholarship
to AMJ; the Instituto Internacional de Educagao do Brasil (IEB) and the Gordon and
Betty Moore Foundation for the Scholarship of Studies on Amazonia Conservation
(BECA) to AMJ and TG.
Literature cited
Coelho G. 2008a. Diplomitoporus taquarae. Fungal Planet 25.
Coelho G. 2008b. Diplomitoporus marianoi-rochae. Fungal Planet 26.
Domanski S. 1970. Wood-inhabiting fungi of Bialowieza virgin forests in Poland. XIII. Two species
of Diplomitoporus Doman., gen. nov. Acta Societatis Botanicorum Poloniae 39: 191-207.
Gibertoni TB, Ryvarden L, Cavalcanti MAQ. 2004. Studies in neotropical polypores 18. New species
(Basidiomycota) from Brazil. Synopsis Fungorum 18: 44-56.
Lisboa PLB. 1997. (Org.). Caxiuana. Belém (Brazil). 440 pp.
Lisboa PLB. 2002. (Org.). Caxiuana: Populac6ées tradicionais, meio fisico e diversidade bioldgica.
Belém (Brazil). 738 pp.
Loguercio-Leite C, Wright JE. 1998. Diplomitoporus dilutabilis a new species of Polyporaceae
(Aphyllophorales) from Santa Catarina Island, Brazil. Mycotaxon 68: 47-51.
Ryvarden L, Gilbertson RL. 1993. European Polypores. Vol. 1. Fungiflora: Oslo (Norway). 387 pp.
Ryvarden L, Iturriaga T. 2003. Studies in neotropical polypores 10. New polypores from Venezuela.
Mycologia 95(6): 1066-1077.
Ryvarden L. 1991. Genera of Polypores. Nomenclature and taxonomy. Fungiflora: Oslo (Norway).
226 pp.
Sotao HMP, Gugliotta AM, Oliveira A, Luz AB, Melo OA. 2002. Fungos Polipordides. Pp. 433-444.
In: Lisboa PLB (Org.). Caxiuana: Populacées tradicionais, meio fisico e diversidade bioldgica.
Belém (Brazil).
198 ... Martins-Junior, Gibertoni & Sotao
Sotao HMP, Hennen JE, Gugliotta AM, Melo AO, Campos EL. 1997. Fungos - Basidiomycotina. Pp.
213-219. In: Lisboa PLB (Org.). Caxiuana. Belém (Brazil).
Watling R. 1969. Colour Identification Chart. Her Majesty's Stationary Office, Edinburgh
(Scotland).
.
MYCOTAXON
Volume 106, pp. 199-202 October-December 2008
Paradoxa gigantospora comb. nov. from China
YUN WANG? & HuNG-TaAo Hv?
wangy@Crp.cri.nz
' New Zealand Institute for Crop & Food Research Limited
Private Bag 4704, Christchurch, New Zealand
*Department of Forestry, National Taiwan University
106 Taipei, Taiwan
Abstract — A new combination, Paradoxa gigantospora, is proposed for the truffle Tuber
gigantosporum that was first described in 1991 based on a specimen collected from
southwestern China. Paradoxa gigantospora is distinct in that it invariably has only
one spore per ascus, a defining character for the genus Paradoxa. This represents the
second species for the genus Paradoxa and the first time this genus has been recorded
in China.
Keywords — Pezizales, taxonomy, fungus
Introduction
In 1991, anew truffle species, Tuber gigantosporum was described by Wang & Li
(1991) based on a specimen collected in 1988 from Huidong County, Sichuan,
China. The species was distinguished by ascospores larger than those produced
by all other species of Tuber. The species also had the distinction of having only
invariably one ascospore per ascus. A second collection of Tuber gigantosporum
was obtained from Yongren County, Yunnan, China in 2002.
In 1935 Mattirolo established the genus Paradoxa based on Paradoxa monospora
Mattir., which is characterized by its invariably 1-spored asci as illustrated by
Montecchi & Sarasini (2000). We now believe that Tuber gigantosporum is
congeneric with Paradoxa Mattir. and here transfer the species from Tuber to
| Paradoxa.
Materials and methods
| The type specimen of Tuber gigantosporum was collected from Huidong County,
Sichuan, China in 1988 and the second specimen from Yongren County,
Yunnan, China in 2002. They are deposited in the Herbarium of Institute of
200 ...Wang & Hu
Applied Ecology, Chinese Academy of Sciences (IFS, not yet listed in the Index
Herbariorum). Macroscopic characters were described from fresh specimens.
Photographs were made and microscopic features described after examining
fresh material or freeze microtome sections in cotton blue under a Nikon E400
microscope.
Taxonomy
Paradoxa gigantospora (Y. Wang & Z. P. Li) Y. Wang comb. nov. FiGuREs 1-6
MycoBANK MB 512137
BasionymM: Tuber gigantosporum Y. Wang & Z. P. Li, Acta Mycol. Sinica 10(4): 263.
Loot
EryMo.oey: the specific epithet refers to the giant ascospores of this species.
AScoMATA (Fig. 1) solid, irregularly subglobose and slightly lobed, pale
yellow-brown to brown, up to 15 mm broad, rough with yellow-brownish,
scurfy patches and lines. ODor not distinctive. PERIDIUM (Fig. 2) 190-230 um
thick, of interwoven hyphae 3-4 um broad at the septa, many cells inflated .
up to 10 um, the out-layer 70-90 um thick and brownish, the inner-layer
hyaline and grading into the gleba tissue. GLEBA (Fig. 3, 4) whitish in youth,
becoming brown to dark reddish brown at maturity, marbled with distinct,
whitish, branched veins merging with the peridium at many points and some
penetrating the peridium to the ascomatal surface; fertile tissue of hyaline,
interwoven hyphae, 3-4 um broad at the septa, many cells inflated up to 20 um.
Asct invariably 1-spored, mostly subglobose, stemless, prior to spore formation
hyaline, thin-walled, subglobose, when containing a spore hyaline, ellipsoid,
160 x 140 um, the wall up to 4 um thick, randomly enclosed in the fertile tissue.
Ascosporks (Fig. 5 and 6) pale brown in youth, becoming dark reddish brown
at maturity, ellipsoid, (80-)105-115(-120) x (55-)70-75(-80) um excluding
ornamentation of a reticulum up to 2 um tall, the meshes irregularly polygonal,
varying in size, 7-8 along the spore and 3-5 across.
ECOLOGY AND DISTRIBUTION — Hypogeous under Pinus yunnanensis and
Quercus spp., August to December. Known from Huidong County, Sichuan
and Yongren County, Yunnan, China.
SPECIMENS EXAMINED — CHINA. Sichuan province: Huidong County, August 1988, Y.
Wang 89921 (IFS 89921 - holotype). Yunnan: Yongren County, 17 December, 2002, H.
T. Hu (IFS 0201).
COMMENTS - Mattirolo (1935) proposed the genus Paradoxa for P. monospora,
which is distinguished by having only one spore per ascus (Montecchi &
Sarasini 2000). It is, therefore, appropriate to transfer Tuber gigantosporum from
Tuber to Paradoxa. The two Paradoxa species are closely related by their similar
ascomata, gleba and spore color in addition to the one-spored asci. However,
P. gigantospora differs from P monospora in having very large, ellipsoid
Paradoxa gigantospora comb. nov. (China) ... 201
Figs. 1-6. Fig. 1. Paradoxa gigantospora. Section of ascorcarp showing the gleba, Hu 0201 (bar = 2.0
mm). Fig. 2. Section of peridium, Wang 89921 (bar = 60 um). Fig. 3. Section of gleba showing the
ascal tissues, composed of 1 spored-asci with no stalks, Wang 89921 (bar = 250 um). Fig. 4. Section
of gleba showing the interascal tissues, composed of colourless, interwoven hyphae 3 - 4 um in
diameter, Wang 89921 (bar = 60 um). Fig. 5. Asci and ascospores, Wang 89921 (bar = 90 um).
Fig. 6. Ascospore, Wang 89921 (bar = 40 um).
ascospores (105-115 x 70-75 um) compared with those of P monospora
that measure 50 x 60 (- 70) um. This is only the second species of the genus
Paradoxa and the first time this genus has been recognized in China.
These two closely related species are widely separated: P monospora is
known only from Italy and P. gigantospora only from southwestern China.
This implies a close biogeographical relationship between the Mediterranean
and southwestern China. This hypothesis is also supported by the results of
studies of Chinese truffle flora in the southwestern China over the past 20 years
202 ...Wang & Hu
that show interesting parallels between the two regions. For example, Tuber
uncinatum (=Tuber aestivum) and Tuber excavatum were found in both regions
(Song et al. 2004, Wang at al. 2008). Likewise, Chinese black truffles, the Tuber
indicum complex in southwestern China, resemble the European Périgord
black truffle (Tuber melanosporum) (Yamanaka et al. 2001, Jeandroz at al. 2008).
Finally, Tuber pseudoexcavatum from southwestern China is closely related to
the European species Tuber brumale (Jeandroz at al. 2008).
Acknowledgments
We appreciate the valuable review comments of Drs. J. Trappe, I.R. Hall and A.
Zambonelli. We also thank Nina Hasom-Williams for correcting the English. This study
was supported by Kunming Institute of Botany and Institute of Applied Ecology Chinese
Academy of Sciences, China.
Literature cited
Jeandroz S, Murat C, Wang YJ, Bonfante P, Tacon FL. 2008. Molecular phylogeny and historical
biogeography of the genus Tuber, the “true truffles”. Journal of Biogeography 35: 815-829.
Mattirolo O. 1935. Catalogo ragionato dei funghi ipogei raccolti nel Canton Ticino e province
italiane confinanti. BeitrageSchweiz 8(2): 1 — 59.
Montecchi A, Sarasini M. 2000. Fungi ipogei d’Europa. Fondazione Centro Studi Micologici DELLs
A.M.B. Vicenza. pp. 244-246.
Song MY. 2004. Tuber huidongense sp. nov. from China. Mycotaxon 83 191-194.
Wang Y, Li ZP. 1991. New species of Tuber from China. Acta Mycologica Sinica 10(4): 263-265.
Wang Y, Peigui L, Chen J, Hu HT. 2008. China - a newly emerging truffle-producing nation. In:
Actes du Collogue: La Culture de la truffe dans le monde, Brive-la-Gallarde — 2 Février 2007
(ed. Schvalier, G. & Raynal B.) - Le Causse Corrézien: Brive, France. pp. 35-44.
Yamanaka K, Namba K, Tajiri A. 2001. Asian black truffles. In: Science et Culture de la Truffe. Actes
du V° Congrés International, 4 au 6 mars 1999, Aix-en-Provence, France. pp. 117 -120.
MY COTA XON
Volume 106, pp. 203-207 October-December 2008
A new species of Phaeoramularia on Cimicifuga (Ranunculaceae)
FENG-YAN ZHAI* YING-LAN GUO? YING-JIE Liu? Yu Li?
' fengyan780103@163.com | rabbitying@163.com
Henan Institute of Science and Technology
Henan 453003, China
? guoyl@sun.im.ac.cn
Institute of Microbiology, Chinese Academy of Science
Beijing 100101, China
** yuli966@126.com
JiLin Agricultural University
ChangChun 130118, China
Abstract —A new species, Phaeoramularia cimicifugae on Cimicifuga dahurica
(Ranunculaceae) is reported. Latin diagnosis, illustration and English description of the
new species are provided. Distinctions between the new species and its closely related
species — P. clematidis, P. lomaensis, P. sudanensis and P. delphinii — are discussed. The
type and isotype specimens are deposited in HMAS and HMJAU.
Key words —imperfect fungi, taxonomy
Introduction
A fungus occurring on Cimicifuga specimens was collected from Inner
Mongolia in China. Its multiseptate, cylindrical, catenate conidia produced
on fasciculate conidiophores that emerged through stomata indicated that it
_ belonged in the genus Phaeoramularia. After comparing our specimen with all
— other Phaeoramularia species previously reported on Ranunculaceae, we found
it different enough to propose as a new Phaeoramularia species, described
below.
Taxonomy
Phaeoramularia cimicifugae F.Y. Zhai, Y.L. Guo & Yu Li, sp. nov. FiG.1
MycoBAnk MB 512082
Maculae amphigenae, punctatae, orbiculares vel irregulares, 3.0-10.0 mm diam.,
aliquando confluentes, atro-brunneae, centro luteo-brunneae, extus halonatis luteis vel
luteo-brunneis. Caespituli amphigeni. Mycelium immersum. Stromata nulla vel parva.
*Corresponding author
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Phaeoramularia cimicifugae sp. nov. (China) ... 205
20m
Fig.1 Phaeoramularia cimicifugae
Conidiophores and conidia (HMAS 143916)
Conidiophora emergena per stomata, laxe vel dense fasciculata, olivaceo-brunnea, laevia,
ramosa, recta vel curvata, 0-7-geniculata, ad apicem truncata vel conico-truncata, 0-3-
septata, 25.0-91.0 x 3.9-7.8 um. Cicatrices conspicue incrassatae, 2.0-3.0 um latae.
Conidia cylindrica, pallide olivaceo-brunnea vel olivacea brunnea, catenata, laevia, recta
vel leviter curvata, ad apicem conico-truncata, ad basin obconico-truncata, 1-5-septata,
imprimis 1-3-septata, 23.0-57.0 x 2.6-5.2 um.
Hab. in foliis vivis Cimicifugae dahuricae (Turcz.) Maxim. (Ranunculaceae), Motianling,
Xingan, Inner Mongolia, 9 VII 1991, leg. Y.L. Guo, no. 1646 (HMAS 143916, Holotypus:
HMJAU 30004, Isotypus), no. 1658 (HMAS 143917).
Leaf spots amphigenous, punctiform, circular or irregular, 3.0-10.0 mm diam.,
sometimes confluent, blackish brown, with a yellowish brown center and yellow
to yellowish brown halo on the upper surface, paler on the lower surface. Fruiting
amphigenous. Mycelium immersed. Stromata none or small. Conidiophores
emerging through stomata, loosely or densely fasciculate, olivaceous brown,
paler towards the apex, smooth, branched, straight to curved, 0-7-geniculate,
regular or irregular in width, sometimes narrower towards the apex, truncate
or conically truncate at the apex, 0-3-septate, 25.0-91.0 x 3.9-7.8 um. Conidial
scars conspicuously thickened, 2.0-3.0 um wide. Conidia cylindrical, pale
olivaceous brown to olivaceous brown, catenulate, smooth, straight or slightly
curved, conically truncate at the apex, obconically truncate at the base, 1-5-
septate, chiefly 1-3-septate, 23.0-57.0 x 2.6-5.2 um.
CoMMENTs: Crous & Braun (2003) reduced Phaeoramularia to synonymy
with Passalora, which mainly differs from Phaeoramularia by forming
206 ... Zhai & al.
solitary conidia. Crous & Braun (2003) maintain that amongst cercosporoid
hyphomycetes the formation of single or catenate conidia was not tenable
as a distinguishing generic character, citing support from ITS and 5.8S
rDNA sequence analysis (Crous et al. 2001). However, we wish to emphasize
morphological characteristics and believe that the mode of conidia formation,
singly or in chains, is a major character delimiting cercosporoid hyphomycetes,
including Phaeoramularia and Passalora. We therefore prefer to recognize
Phaeoramularia as a separate genus.
Four other Phaeoramularia species also occur on Ranunculaceae: P. clematidis
S.K. Singh & R.K. Chaudhary (Singh et al. 1995), P lomaensis Deighton
(Deighton 1979), P sudanensis Deighton (Deighton 1979) and P delphinii FY.
Zhai et al. (Zhai et al. 2007). Among the four species, however, P. clematidis,
P. lomaensis and P. sudanensis are associated with Clematis, while P. delphinii is
associated with Delphinium; none of them is associated with Cimicifuga.
Phaeoramularia clematidis (found on Clematis gouriana Roxb.) exhibits
hypogenous fruiting, well-developed stromata, bears 5-8 conidiophores in a
fascicle that are unbranched, darker, shorter and narrower and bear smaller
conidial scars than found in P. cimicifugae, and produces conidia that are mostly
solitary, occasionally catenate, or (more rarely) branched, subcylindrical, paler,
1-2-septate, and shorter.
P. lomaensis and P. sudanensis (also on Clematis spp.) do not form leaf spots,
exhibit hypogenous fruiting, bear ~12 conidiophores per fascicle that are paler,
usually branched, rarely simple, sinuous or geniculate, longer, narrower and
bear smaller conidial scars, and produce conidia that are paler, longer, and
slightly wider. In addition P lomaensis forms well-developed stromata and
subcylindric conidia, and P. sudanensis produces slightly curved or sometimes
slightly sigmoid conidia.
P. delphinii (found on Delphinium sp.) possesses well-developed stromata,
produces conidiophores that are fairly densely fasciculate, unbranched, paler,
shorter and narrower and bear smaller conidial scars, and produces conidia
that are catenulate and often in branched chains, ellipsoid to cylindric, fewer
septate, paler, shorter and wider.
Acknowledgements
We express our deep appreciation to Prof. John Webster and Jian-Yun Zhuang for
their valuable suggestions and earnest assistance in reviewing earlier drafts of this
manuscript.
Phaeoramularia cimicifugae sp. nov. (China) ... 207
Literature cited
Crous PW, Braun U. 2003. Mycosphaerella and its anamorphs: 1. Names published in Cercospora
and Passalora. CBS Biodiversity Series 1: 19-22
Crous PW, Kang JC, Braun U. 2001. A phylogenetic redefinition of anamorph genera in
Mycosphaerella based on ITS rDNA sequence and morphology. Mycologia 93(6): 1081-1101.
Singh SK, Chaudhary RK, Morgan-Jones G. 1995. Notes on Hyphomycetes. LXVII. Three new
species of Phaeoramularia from Nepal. Mycotaxon 54: 57-66
Deighton FC. 1979. Studies on Cercospora and allied genera. VII. New species and redispositions.
Mycol. Pap. 144: 32-45
Zhai FY, Guo YL, Li Y. 2007. A new species of Phaeoramularia on Ranunculaceae. Mycotaxon 100:
19-192
’
Volume 106, pp. 209-217 October-December 2008
Hyaloscyphaceae (Ascomycota)
growing on tree ferns in Mexico
ROSARIO MEDEL’& FRANCISCO LOREA-HERNANDEZ?”
rosario.medel@inecol.edu.mx & francisco.lorea@inecol.edu.mx
‘Instituto de Ecologia, Unidad de Micologia,’ Depto. Biologia Evolutiva
Apartado Postal 63, Xalapa, 91000, Veracruz, Mexico
Abstract — Five species of Lachnum growing on fern leaves in cloud forest are reported
from Mexico. Lachnum fimbriiferum, L. oncospermatis and L. singerianum are new
records for the country, L. pteridophyllum and L. varians are new records for Veracruz
State.
Key words — ascomycete, discomycete, diversity
Introduction
| Hyaloscyphaceae is one of the major families in the Leotiomycetes, with more
_ than 500 species (Kirk et al. 2001) and a worldwide distribution. In Mexico this
- family has not yet been monographed and few references about this group of
_ fungi are available (Chacén & Guzman 1983, Chacon & Medel 1993, Galan et al.
1994, Medel & Calonge 2004). The authors found several ascomycetes growing
| on tree fern leaves while studying the diversity of Ascomycota in cloud forest in
_ the Veracruz State. A literature search showed that only two species, Lachnum
varians (as D. varians var. varians) from Chiapas and Lachnum pteridophyllum
| (as D. varians var. pteridophyllum) from Oaxaca, have been reported from
_ Mexico growing on this type of host (Cantrell 2004, Haines 1980, Nagao 1996,
| Nagao & Doi 1996, Samuels & Rogerson 1990, Raitviir 2004, Spooner 1987, Wu
et al. 1998, Zhuang & Wang 1998). The only checklist of fungi from the cloud
forest (Chacon & Medel 1993) cites 594 species of fungi; of the 96 ascomycetes
_ listed, only one species, Lachnum brasiliense (Mont.) J.H. Haines & Dumont
| from Veracruz, belongs to the Hyaloscyphaceae. Tree ferns in Mexican cloud
_ forest are one of the most conspicuous and abundant elements, easily seen
| because of their shape and size. The aim of this study is to report the species of
| Hyaloscyphaceae associated with tree ferns, especially those found in the cloud
forests of Veracruz, Mexico.
210 ...Medel & Lorea-Hernandez
Materials and methods
Specimens were collected in 1986-89 and 2004-06 during two surveys of of
cloud forest remnants in the central region of Veracruz. Tissues were examined
microscopically in 5% KOH to record measurements or in Melzer’s reagent
(MLZ) to study the amyloid reaction; tissues from one species were mounted
in Lugol solution (IKI) to determine hemiamyloidity. Spore averages (denoted
by x) were determined from 25 spores per species; Q-values are cited according
to Largent et al. 1977. All studied specimens are kept in the Mycological
Collection of the Instituto de Ecologia herbarium (XAL). Plant hosts were
named to species when samples were suitable for reliable identifications.
Studied species
Lachnum fimbriiferum (Berk. & M.A. Curtis) J.-H. Haines,
Nova Hedwigia 54: 103 (1992) Fics 1-2, 11-12
= Peziza fimbriifera Berk. & M.A. Curtis in Berkeley, J. Linn. Soc., Bot. 10: 367 (1868)
= Dasyscyphus fimbriifer (Berk. & M.A. Curtis) Sacc., Syll. Fung. 8: 452 (1889)
Apothecia globose to cup shaped, 0.3-0.5 mm diam, disc pale yellow or cream
color when dry, covered with superficial hairs white to yellowish, stipitate. Asci
70-80 x 8-9 um, in KOH, cylindrical, MLZ+, with a small blue Hymenoscyphus-
type plug, 8-spored, without croziers. Spores fusiform with rounded ends,
(15—) 20-30 x 3 (4) um, x = 11.2 x 1.2 um (Q>3.0), slightly curved, with small
globules in MLZ, KOH. Paraphyses filiform, longer than asci up to 2 um, 1-3 um
diam, hyaline, simple to branched and septate near the base. Hairs cylindrical,
100-300 x 4-5 um, septate, hyaline, finely granulate.
HABITAT: gregarious, on decaying leaves of Alsophila firma (Baker) D.S. Conant,
Cyathea bicrenata Liebm., other Cyatheaceae spp. and Dicksonia sellowiana
Hook., in cloud forest at 1400-1680 m altitude.
KNOWN DISTRIBUTION: Colombia, Cuba, Dominican Republic, Ecuador,
Jamaica, Venezuela and Mexico (reported here).
STUDIED MATERIAL: MEXICO. VERACRUZ. Municipality of San Andrés
Tlalnehuayocan, Rancho Agiita Fria, road to San Andrés Tlalnehuayocan, 18/5/2004,
Medel 869; 23/7/2005, Medel 973, 976, 982, 985, 988, 990; Municipality of Rafael Lucio,
Santa Barbara Farm, km 10 NE highway Xalapa-La Joya, 3/10/1986, Medel 200, 209; 2/6/
1987, Medel 360, 361, 366; 14/5/1987, Medel 343, Chacon 1672.
Notrs— ‘This species is not easy to recognize using macroscopic characters
since many species of Lachnum growing on tree ferns are similar, but the cream
color of the disc and the white hairs are diagnostic. Spore, ascus, and paraphysis
sizes differentiate this species from L. singerianum, a related species that
Haines (1980) considered a variety of L. fimbriiferum. In comparing spore sizes,
however, we found enough differences in the studied material to recognize
Hyaloscyphaceae on tree ferns (Mexico) ... 211
Figs. 1-10. 1-2: Lachnum fimbriiferum. 1: asci, paraphyses and ascospores; 2: ascospores close up
(Medel 869). 3-4: L. oncospermatis. 3: asci and paraphyses; 4: ascospores close up (Medel 333);
5-6: L. pteridophyllum. 5: asci: paraphyses and ascospores; 6: asci and ascopores close up (Medel
870); 7: L. singerianum. Asci, ascospores and paraphyses (Medel 923); 8-10 L. varians. 8: asci and
ascospores; 9: ascospores close up; 10: hairs showing amorphous resin masses (Tapia 851).
Scale bar: 10 um; figure 5: 154m. Photographs R. Medel.
212 ...Medel & Lorea-Hernandez
two independent species: spore lengths never exceed 10 um in L. singerianum,
in contrast to the 15-30 um range in L. fimbriiferum. Although this was the
most common species found on ferns in our study, this is the first record for
Mexico.
Lachnum oncospermatis (Berk. & Broome) M.L. Wu & J.H. Haines,
in Wu, et al., Mycotaxon 67: 346 (1998) Fics 3-4, 13
= Peziza oncospermatis Berk. & Broome, J. Linn. Soc. Bot. 14: 105 (1873)
= Dasyscyphus oncospermatis (Berk. & Broome) Sacc., Syll. Fung. 8: 452 (1889) .
Apothecia infundibuliform, 0.3-0.5 mm diam, disc dark yellow or stramineus
when dry, covered with buff yellow or whitish hairs, stipitate, some apothecia
have bifurcate stipes. Asci cylindric-clavate, 80-90 x 10-11 um in KOH, with
a small pore, Hymenoscyphus-type, MLZ+, without croziers. Ascospores long
fusiform with one end wider than the other, 30-40(-45) x 2-3 um, x = 35.3 x
2.5 um (Q>3.0) with clearly small globules in KOH. Paraphyses filiform, 1-2
um. Hairs cylindrical, 3-4 um, septate, yellowish, roughened by granules.
HABITAT: gregarious on decaying leaves of Dicksonia sellowiana, in cloud forest
at 1680 m altitude.
KNOWN DISTRIBUTION: Australia, Indonesia, Taiwan, The Philippines, Sri
Lanka, and Mexico.
STUDIED MATERIAL: MEXICO. VERACRUZ. Municipality of Rafael Lucio, Santa
Barbara Farm, km 10 NE highway Xalapa-La Joya, 14/10/1987, Medel 333.
Notes— ‘The solitary or bifurcate stipe in some apothecia and the long spores
are diagnostic for this species. Haines (1980) cites spores up to 32 um for the
typical variety of L. oncospermatis and refers specimens with longer (36-67 x
2.5-3.5 um) spores to L. oncospermatis var. macrospora. Wuetal. (1998) reported
spores of (24—)27-39(-42) x 2-3 um. Spores from Medel 333 are up to 45 um
and agree with the spore size recognized by Wu et al. (1998) for this species.
Lachnum varians, which produces similarly colored ascomata, produces spores
shorter than in L. oncospermatis. Haines (1980) mentioned that this species
grows on Cyatheaceae and Liliaceae, and the record on Oncosperma from
Ceylon is not confirmed. Wu et al. (1998) reported this species growing on two
genera of Cyatheaceae from China. The material from Mexico was growing on
Dicksonia, a taxon not previously known as a host for this species. This is the
first record for Mexico.
Lachnum pteridophyllum (Rodway) Spooner,
Biblioth. Mycol. 116: 470 (1987) Fires 5-6, 14
= Dasyscyphus pteridophyllus Rodway, Pap. Proc. R. Soc. Tasmania 1920: 158 (1921)
= Dasyscyphus varians var. pteridophyllus (Rodway) J.H. Haines, Mycotaxon 11: 209 (1980)
= Lachnum varians var. pteridophyllum (Rodway) M.P. Sharma, Nova Hedwigia 43: 411 (1986)
Hyaloscyphaceae on tree ferns (Mexico) ... 213
Figs. 11-16. 11-12: Lachnum fimbriiferum. 11 apothecia; 12 apothecium close up (Medel 869);
13: L. oncospermatis. Apothecia showing bifurcate stipes (Medel 333); 14: L. pteridophyllum.
Apothecia (Medel 870); 15: L. singerianum. Apothecium (Medel 923); 16: L. varians. Apothecium
(Tapia 851). Scale bar: 0.5 mm. Photographs R. Medel.
214 ...Medel & Lorea-Hernandez
Apothecia globose to funnel shaped, 0.3-0.5 mm diam, disc light to dark yellow
when dry, covered with pale yellow hairs, stipitate. Asci cylindric-clavate,
40-55(-60) x 5-6(-7) um, with a blue pore Hymenoscyphus-type MLZ+, 8-
spored, without croziers. Ascospores narrowly fusiform, 11-15(-17) x 1-2 um,
x = 12.3 x 1.7 um (Q>3.0), with small globules in KOH, less evident in MLZ.
Paraphyses narrow, lanceolate, up to 2 um diam. Hairs 100-200 x 4-5 um,
hyaline to yellow in mass.
HABITAT: gregarious on decaying leaves of Dicksonia sellowiana, Cyathea
divergens var. tuerckheimii (Maxon) R.M. Tryon and other Cyathea spp. At
1400-1680 m altitude.
KNOWN DISTRIBUTION: Colombia, Dominican Republic, Jamaica, Mexico,
Panama, Peru, Puerto Rico, New Guinea, New Zealand, Taiwan, Tasmania and
Venezuela.
STUDIED MATERIAL: MEXICO. VERACRUZ. Municipality ofSan Andrés Tlalnehuayocan,
Rancho Agiiita Fria, road to San Andres Tlalnehuayocan, May 8, 2004, Medel 860, 867,
870; 23/5/2004, Medel 880, 882; July 23, 2005, Medel 978; Municipaliy of Rafael Lucio,
Santa Barbara Farm, km 10 NE highway Xalapa-La Joya, October 14, 1987, Medel 368.
Notes— Lachnum pteridophyllum is distinguished by spores with acute ends
that are not longer than 20 um (Spooner 1987). It is morphologically very similar
to L. varians, which produces shorter (< 15 um) spores with rounded ends.
Haines (1980) treated L. pteridophyllum as a variety of L. varians, but Spooner
(1987) later argued that spore size and shape differences justified elevation of
the taxon to species level. Haines (1980) first reported L. pteridophyllum from
Oaxaca State. This is the first record for Veracruz State.
Lachnum singerianum (Dennis) W.Y. Zhuang & Zheng Wang,
Mycotaxon 67: 27 (1998) Fics 7, 15
= Dasyscyphus singerianus Dennis, Kew Bull. 13: 465 (1959 [‘1958’])
= Dasyscyphus fimbriifer var. singerianus (Dennis) J. H. Haines, Mycotaxon 11: 199
(1980) .
Apothecia discoid to globose, 0.3-0.5 diam., disc whitish yellow or pale orange
when dry, covered with whitish to yellow hairs, shortly stipitate. Asci cylindric-
clavate 45-55 x 5-6 um, with a blue pore Hymenoscyphus-type, MLZ+, without
croziers. Ascospores ellipsoid, (7-)8-10 x 1(-2) um, x = 9.2 x 1.7 um (Q>3.0),
with evident globules in KOH, non-septate, hyaline. Paraphyses cylindric up to
2 um diam, hairs cylindrical, 150-200 x 4-5 um, septate, finely granulate.
HABITAT: gregarious, on decaying leaves of Alsophila firma and Dicksonia
sellowiana at 1400-1680 m altitude.
KNOWN DISTRIBUTION: in the Andean region of Bolivia to Venezuela, China,
Jamaica, and Mexico.
Hyaloscyphaceae on tree ferns (Mexico) ... 215
Studied material: MEXICO. VERACRUZ. Municipality of Xalapa, Botanical Garden
Francisco J. Clavijero, km 2.5 old highway Xalapa-Coatepec, August 3, 1989, Medel 445
(XAL); municipality of San Andrés Tlalnehuayocan, road to San Andres Tlanehuayocan,
Rancho Aguita Fria, June 6, 2005, Medel 923; October 12, 2005, Medel 1122.
Notes— Haines (1980) first cited L. singerianum as a variety of Dasyscyphus
fimbriifer, but Zhuang and Wang (1998) regard it as an independent species.
They cite a wider (< 3 um) spore size than found in the Veracruz specimens
(Zhuang & Wang 1998), which otherwise have the general morphology
and host (Cyatheaceae, cf. Haines 1980). Lachnum singerianum resembles
L. fimbriiferum, which differs in its longer ascospores, asci, and paraphyses.
This is the first report of L. singerianum from Mexico.
Lachnum varians (Rehm) M.P. Sharma, Nova Hedwigia 43: 411 (1986) Fras 8-10, 16
= Dasyscypha varians Rehm, Hedwigia 39: 94 (1900)
Apothecia globose to funnel shaped 0.3-0.5 mm diam, disc pale orange when
dry, stipitate, densely covered with whitish or pale buff hairs with amorphous
masses of resin, which do not dissolve in KOH or Melzer’s reagent. Ascospores
long elliptical with rounded ends, 10-15 x 3-4(-5) um, x = 12.9 x 4.1 um
(Q>3.0). Asci cylindric-clavate, 70-80 x 10 um, MLZ-, hemiamyloid in Lugol's
solution, without croziers. Paraphyses filiform up to 2 um diam. Hairs cylindric,
up to 100 x 3-5 um, septate, thin-walled.
HABITAT: gregarious on decaying leaves of Alsophila firma in cloud forest at
1300-1680 m altitude.
KNOWN DISTRIBUTION: Australia, Hawaii, New Guinea, New Zealand, South
America, The Caribbean, and Mexico.
STUDIED MATERIAL: MEXICO. VERACRUZ. Municipality of Xalapa, around the
Conecalli, km 2.5 old higway Xalapa-Coatepec, August 27, 1991, Tapia 851; municipality
of San Andres Tlanehuayocan, road to San Andrés Tlalnehuayocan, Rancho Agitita Fria,
May 23, 2004, Medel 882.
Notes—Lachnum varians is characterized by short (< 15 tm) spores with
rounded ends and MLZ- asci. Although spores of studied material were wider
than those referred by Haines (1980) and Spooner (1987) for the species, other
characters fit the concept of L. varians including the host (Cyatheaceae family).
It is close to L. pteridophyllum, which differs in its longer spores with acute
ends. Haines (1980) previously reported L. varians for Mexico from Chiapas
State, but this is a first record from Veracruz State.
Discussion
Tree ferns are common to Mexican cloud forests, and there is a great diversity
of fungi associated with these plants. The five Lachnum species described
216 ...Medel & Lorea-Hernandez
above were the most common ascomycetes growing on Mexican tree ferns,
but there are-other small unidentified discomycetes (e.g. species of Orbilia Fr.
and Cyathicula De Not.) represented in the cloud forests as well. Sixty percent
of eight taxa cited by Haines (1980) known to occur on tropical ferns are
represented in Mexico. The fragmentary distribution and extensive human use
of the Mexican cloud forests make them vulnerable and likely to disappear.
There is a decline on the diversity of tree ferns in Mexico and the associated
fungi are endangered.
Acknowledgements
Thanks to Dr. Donald Pfister (Farlow Herbarium, Harvard University) who critically
read the first version of this work. Especially thanks to the reviewers: Dr. Sharon
Cantrell (Universidad de Tubago, Puerto Rico) and Dr. Seppo Huhtinen (University
of Turku, Finland) for critical review this paper and for all observations to improve
the manuscript. Thanks to the following technicians of Instituto de Ecologia: Juan Lara
Carmona helped on herbarium work, Claudia Gallardo collected some fern samples for
identification, Maricruz Peredo edited photographs and Manuel Hernadez help in the
edition of the plates. |
Literature cited
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Republic and the Caribbean region. Caribbean Journal of Science 40: 139-144.
Chacon S, Guzman G. 1983. Ascomycetes poco conocidos en México. Boletin de la Sociedad
Mexicana de Micologia 18: 183-218.
Chacon S, Medel R. 1993. Los hongos principalmente macromicetos registrados del bosque
mesofilo de montafia de México. Reporte Cientifico especial, Facultad de Ciencias Forestales
UANL No. 13: 61-110.
Dennis RWG. 1954. Some inoperculate discomycetes of tropical America. Kew Bull. 2: 289-348.
Dennis RWG. 1979. Fungus Flora of Venezuela and adjacent countries. Cramer, Lehere.
Galan R, Raitviir A, Ayala N, Ochoa C. 1994. First contribution of the knowledge of the Leotiales of
Baja California and adjacent areas. Mycological Research 98:1137-1152.
Haines JH. 1980. Studies in the Hyaloscyphaceae I: some species of Dasysycphus on tropical ferns.
Mycotaxon 11: 189-216.
Kirk P, Cannon PF, David JC, Stalpers JA. 2001. Ainsworth & Bisby’s Dictionary of the fungi. 9 ed.
CABI Bioscience, Bakeham Lane UK.
Largent D, Johnson D, Watling R. 1977. How identified mushroom to genus III. Microscopic
features, Mad Rivers, Eureka, Ca.
Medel R, Calonge FD. 2004. Aportacion al conocimiento de los discomicetes de México, con especial
referencia al género Helvella. Boletin de la Sociedad Micologica de Madrid 28:151-159.
Nagao H. 1996. Discomycetes on decayed tree ferns: (2). Lachnum varians (Rehm) Spooner and
Lachnum sclerotii (A.L. Smith) Haines et Dumont new to Japan. Bulletin of the National Science
Museum Series B 22(3): 105-111.
Nagao H, Doi Y. 1996. Discomycetes on decayed tree ferns. (1) Lachnum pteridophyllum (Rodway)
Spooner New to Japan. Bulletin of the National Science Museum Series B 22(1): 19-22.
Hyaloscyphaceae on tree ferns (Mexico) ... 217
Raitviir A. 2004. Revised synopsis of the Hyaloscyphaceae. Estonian Agricultural University.
Institute of Zoology and Botany, Tartu.
Samuels G, Rogerson CT. 1990. Some Ascomycetes (fungi) occurring on tropical ferns. Brittonia
42: 105-115.
Spooner BM. 1987. Helotiales of Australasia: Geoglossaceae, Orbiliaceae, Sclerotiniaceae,
Hyaloscyphaceae. Biblioteca Mycologica 116, J Cramer, Sttutgart.
Wu ML, Haines JH, Wang YZ. 1998. New species and records of Lachnum trom Taiwan. Mycotaxon
67: 341-352.
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Mycotaxon 67: 21-31.
Volume 106, pp. 219-226 October-December 2008
Rust fungi on Saccharum species from Pakistan
S. H. IQBAL, A. N. KHALID, N. S. AFSHAN & A. R. NIAZI
pakrust@gmail.com
Department of Botany, University of the Punjab
Quaid-e-Azam Campus, Lahore, 54590, Pakistan
Abstract — Four species of rust fungi are described on three species of Saccharum.
Among these, Puccinia melanocephala and P. miscanthi are new records for Pakistan.
Puccinia coronata var. avenae and P. kuehnii are already known from this area. Saccharum
is a new host for the Puccinia coronata species complex from Pakistan.
Key words — Andropogoneae, Poaceae, sugarcane
Introduction
The plant genus Saccharum is placed in the tribe Andropogoneae of family
Poaceae. It is a genus of 35-40 species, occurring throughout the tropics and
extending to warm temperate regions. Nine species of Saccharum occur in
Pakistan and one of them, Saccharum officinarum L. (sugarcane), is cultivated
as an important commercial crop (Cope 1982). Sugarcane, the source of sugar, is
being grown on 0.96 million hectares with a production of 42 million tonnes in
Pakistan. Unfortunately, its full production is hindered due to biotic and abiotic
stresses. Diseases such as red rot, Pokka boeng, smut, rusts, mosaic and red
stripe are some of the major biotic stresses that may reduce the yield by 10-77%
(Ali 2006). Common species of rust fungi that attack Saccharum worldwide
include orange rust, Puccinia kuehnii, PR. melanocephala (= P. erianthi), and
P. miscanthi.
During a survey of rust fungi of Pakistan, it was found that Saccharum is
a favorable host for rust fungi. Four species of rust fungi are herein described
on three species of Saccharum from different localities. Among these, Puccinia
melanocephala and P. miscanthi are new records for Pakistan. Puccinia kuehnii
and P. coronata var. avenae have already been reported from Pakistan, but
Saccharum is a new host for the Puccinia coronata species complex from
Pakistan.
Materials and methods
Specimens were collected from different areas of Pakistan. Dried herbarium specimens
identified as Puccinia kuehnii were borrowed from the Herbarium, Department of
220 ... Iqbal & al.
Botany, University of the Punjab, Lahore (LAH). Freehand sections of infected tissue
and spores were mounted in lactophenol and gently heated to boiling. The preparations
were observed under a NIKON YS 100 microscope and photographed with a digipro-
Labomed. Drawings of spores and paraphyses were made using a Camera Lucida (Ernst
Leitz Wetzlar, Germany). Spore dimensions were measured using an ocular micrometer.
At least 25 spores were measured for each spore stage. The rust fungi were identified by
comparing their characters with the descriptions and illustration made by Cummins
(1971) and Hiratsuka et al. (1992). The rusted specimens have been deposited in LAH.
Description of taxa
1. Puccinia coronata var. avenae W.P. Fraser & Ledingham, Sci. Agric. 13: 315
(1933) (Frcs. AB)
=Puccinia coronifera Kleb., Z. Pflanzenkrankh. Pflanzenschutz 4: 135 (1894).
Spermogonia and aecia unknown in Pakistan. Uredinia on adaxial leaf surface,
mostly amphigenous, cinnamon brown, 0.09-0.2 x 0.2-0.3 mm. Urediniospores
ovoid to obovoid or ellipsoid, 16-21 x 20-31 um; wall 1.5-2 um thick, golden
brown to cinnamon brown, echinulate; germ pores (5-) 9-11, scattered, with ©
slight or no cuticular caps; pedicel hyaline, short, 4-8 x 10-12 um. Paraphyses
clavate to cylindrical, hyaline, 12-19 um thick at apex, 55-60 um long. Telia
abaxial, covered by epidermis, with a few brown stromatic paraphyses, 0.06-0.1
x 0.1-0.2 mm, black. Teliospores clavate to ellipsoid or oblong, not or slightly
constricted at septa, 11-16 x 45-60 (-65) um excluding digitations, digitations
2-several, 6-15 um long; wall 1-1.5 um thick, smooth, chestnut brown, paler
basally; germ pore 1 per cell; pedicel short, not collapsing, brown, 7-9 x 12 um
long.
MATERIAL EXAMINED: On Saccharum ravennae (L.) L., Pakistan, North West Frontier
Province (NWFP), Sheran, 13 August, 2007, NSA # 40 (LAH Herbarium No. NSA
1013).
Puccinia coronata var. avenae was previously reported on Avena and other
grasses (Cummins 1971, Hiratsuka et al. 1992), and on Elymus repens (L.)
Gould (Afshan et al. 2008). According to Cummins (1971), variety avenae has
been proved capable of infecting various grasses, occasionally on grasses where
oats are common. Saccharum is a new host for the Puccinia coronata species
complex from Pakistan.
2. Puccinia kuehnii E.J. Butler, Ann. Mycol. 12: 82 (1914). (PrestC2L
= Uromyces kuehnii W. Kriiger, Ber. Versuchs Stat. f. Zuckerrohr West-Java, Kagok-Tegal 1:
120, 1890. (based on uredinia)
= Uredo kuehnii (W. Kriiger) Wakker & Went, De Ziekten van het Suikerriet Java, Leiden,
p. 144. 1898.
Spermogonia and aecia not found. Uredinia mainly on abaxial leaf surface,
sometimes amphigenous, cinnamon brown to golden brown, 0.09-0.2 x 0.2-0.3
Rusts on Saccharum (Pakistan) ... 221
Figs. A-B: Puccinia coronata var. avenae on Saccharum ravennae
(A). Echinulated urediniospores and clavate paraphyses
(B) Lucida drawing of coronate teliospores.
Scale bars = 10 ttm.
mm. Urediniospores ovoid to obovoid or pyriform, sometimes broadly ellipsoid,
23-30 x 33-48 um; wall up to 3 um thick at sides, sometimes uniformly thick,
but usually with apex up to 9 um thick, golden brown to yellowish brown with
orange granules, moderately echinulate with evenly distributed spines; germ
pores 4-5, equatorial, or tending to equatorial; pedicel hyaline, 6-8 x 24 um.
Paraphyses inconspicuous, not always observed; if present, basal or peripheral,
pyriform to clavate, extremely thin walled, delicate, hyaline to pale brown.
Telia hypophyllous, erumpent, 0.2-0.4 x 0.1-0.5 mm, with few paraphyses.
222 ... Iqbal & al.
ee
ae all
ES
RR,
SF.
Fies. C-D: Puccinia kuehnii
(C). Urediniospores (D). Teliospores.
Scale bars = 10 um.
Teliospores oblong or clavate to fusiform, not or slightly constricted at septa,
18-26 x (26—) 30-40 (-45) um; wall 0.5-1 um thick, smooth, chestnut brown,
paler basally; germ pores obscure; pedicel short, hyaline, 7-8 x 12 um long.
MATERIAL EXAMINED: On Saccharum spontaneum L., S. munja Roxb., S. bengalense
Retz., Pakistan, Punjab, Lahore, Minchin Abad, Faisal abad. 23’ December, 2007. # 43
(LAH).
SPECIMEN EXAMINED: On Erianthus ravennae (L.) P. Beauv. (= Saccharum ravennae),
Lahore, 08" January, 1950, S. Ahmad; Saccharum munja, Lahore, 19'" November, 1950,
S. Ahmad, # 3077; Saccharum ravennae, Lahore, 27" April, 1969, S. Ahmad, # 21855
(LAH).
Puccinia kuehnii was first described as Uromyces kuehnii because of the
presence of apically thick-walled urediniospores that were apparently
mistaken as teliospores. It was later renamed as Uredo kuehnii, since the telial
stage was not found and the apically thick-walled spores were proven to be
urediniospores (Virtudazo et al. 2001). Butler (1914) found teliospores of this
fungus on Saccharum spontaneum and named the species as Puccinia kuehnii.
Virtudazo et al. (2001) made a taxonomic revision of Puccinia species viz. —
P. melanocephala and P. kuehnii, causing rust diseases on sugarcane to clarify
their morphological characteristics.
From Pakistan, P kuehnii is previously reported on S. bengalense,
S. officinarum and S. spontaneum from Lahore and Faisalabad (Ahmad et al.
1997).
Rusts on Saccharum (Pakistan) ... 223
ih Cee
Fics. E-F: Puccinia melanocephala on Saccharum spontaneum
(E) Echinulated urediniospores and capitate paraphyses (F) Teliospores.
Scale bars = 10 um.
3. Puccinia melanocephala Syd. & P. Syd., in Sydow et al., Ann. Mycol. 5: 500
(1907). (Fics. E-F)
= Puccinia erianthi Padwick & A. Khan, Mycol. Pap. 10: 10 (1944).
| Spermogonia and aecia not found in Pakistan. Uredinia mainly on abaxial
leaf surface, sometimes amphigenous, cinnamon brown to dark brown,
subepidermal, erumpent, 0.09-0.2 x 0.2-0.3 mm. Urediniospores obovoid
| to ellipsoid, 21-27 x 28-35 (-38) um; wall 2-2.5 um thick, golden brown to
cinnamon brown, densely echinulate; germ pores 4-5, equatorial; pedicel
hyaline, short, 4-5 x 15-20 um. Paraphyses abundant, capitate, golden brown
| to cinnamon brown, 50-100 um long, with head 15-24 um in diam., 6-9 um
_ thick at lower side, with the wall 1.0-3.0 um thick in the stipe and 5-15 um in
the head. Telia abaxial, covered by epidermis, but early exposed, 0.06-0.1 x
224 ... Iqbal & al.
0.1-0.2 mm, black, with long capitate paraphyses. Teliospores mostly clavate
or ellipsoid to broadly ellipsoid, 16-22 x (28-) 32-45 um, slightly constricted
at septa; wall 1-1.5 um thick, smooth, apex chestnut brown to dark brown, but
paler basally; apex 2-5 um thick, rounded; germ pore 1 per cell, in upper cell
apical or sub-apical, in lower cell near the septum; pedicel short, not collapsing,
brown, 8-10 um wide and up to 12 um long. Three-celled spores occasional.
MATERIAL EXAMINED: On Saccharum spontaneum, Pakistan, North West Frontier
Province (NWEP), Bara Gali, at 2393m a. s. |., 06" November, 2007, # 41 (LAH).
Puccinia melanocephala has previously been recorded on Saccharum officinarum,
Erianthus rufipilus Griseb. and E. ravennae (Cummins 1971).
P. melanocephala is reported for the first time from Pakistan
4: Puccinia miscanthi Miura, Flora of Manchuria and East Mongolia 3: 302 (1928).
(Fics. G-H)
= Puccinia eulaliae Barclay, J. Asiatic Soc. Bengal 60: 216 (1891).
= Puccinia miscanthicola F.L. Tai & C.C. Cheo, Bull. Chinese Bot. Soc. 3: 67 (1937).
Spermogonia and aecia unknown in Pakistan. Uredinia epiphyllous, cinnamon
brown, 0.09-0.2 x 0.2-0.3 mm. Urediniospores subglobose to obovoid or
ellipsoid, 22-28 x 25-33 um; wall 2-3 um thick, light brown to cinnamon
brown, echinulate; germ pores 2-5, equatorial; pedicel hyaline, short, 6-8 x
8-10 um. Paraphyses capitate, hyaline or becoming brown with age, 19-24
tuum thick at the apex and 6-8 um thick at lower side, 78-120 um long. Telia
hypophyllous, covered by epidermis, 0.06-0.1 x 0.1-0.2 mm, black, loculate
with a few brown paraphyses surrounding sori. Teliospores oblong to clavate,
11-19 (-24) x (38-) 40-70 (-75) um, not or slightly constricted at septa; wall 1-
1.5 um thick, smooth, brown to cinnamon brown; apex 4-8 um thick, truncate
or rounded, mostly chestnut brown; germ pore 1 per cell, upper sub-—apical,
lower at the equator or near septum; pedicel short, light brown, not collapsing,
thick walled, 6-7 x 4-13 um. Few one-celled spores also observed.
MATERIAL EXAMINED: On Saccharum ravennae, Pakistan, Northern Areas, Hunza, at
2,440 ma.s. |., 13" August, 2007, # 14. (LAH)
Cummins (1971) reported Puccinia miscanthi on Saccharum narenga Wall.
This is the first report of this species from Pakistan.
Key to the Puccinia species on Saccharum in Pakistan:
1 Teliospores with thick, coronate apex having 2 to several digitations, urediniospores _ |
withlo—9. scattered Seri Ores: sae eee tee. seheeee P. coronata var. avenae —
1* Teliospores with apex rounded to truncate, urediniospores with up to 5, equatorial
PerIN POLES Mi Frei tie sel Peppy ATe tas SI SE Ei oho: RY Uden ere eee eer tone y
2 Telia loculate with a few brown paraphyses surrounding the sori. Teliospores mostly
40-70 um long, with thick, truncate to rounded apex ....... Puccinia miscanthi
Rusts on Saccharum (Pakistan) ... 225
ear
LESS ee
Fics. G-H: Puccinia miscanthi on Saccharum ravennae.
(G) Urediniospores showing equatorial germ pores and capitate paraphyses.
(H) 1-2 celled teliospores with paraphyses.
Scale bars = 10 um.
2* Telia not loculate, with capitate paraphyses, apex thick, rounded to conical.
TeliOcpoLecimostiy less (han 40k ON 0 aera am. ee eee nee er ee 8
3 Teliospores 32-48 um long, with abundant, long, capitate paraphyses, apex thick,
| rounded. Uredinia with distinctly capitate paraphyses, urediniospores with
dense echinulation having regularly spaced spines and uniformly thickened
ISPS its eh RCE cee nner Ra nr eine pe Party es Ontos Puccinia melanocephala
| 3* Teliospores with few capitate paraphyses, apex thick, rounded to conical.
Teliospores 30-40 um long. Uredinia with indistinctly capitate paraphyses,
urediniospores with moderate echinulation having evenly distributed spines
and apically or uniformly thickened walls ................... Puccinia kuehnii
226 ... Iqbal & al.
Acknowledgements
We sincerely thank Dr. Shaun Pennycook, Curator, ICMP, Landcare Research, Auckland,
New Zealand for nomenclature review; Dr. Amy Rossman, Systematic Mycology and
Microbiology Laboratory, USDA-ARS, Beltsville and Dr. Patricia E. Crane, Department
of Renewable Resources, University of Alberta, Edmonton, Alberta, for their valuable
suggestions to improve the manuscript and acting as presubmission reviewers.
References
Afshan NS, Khalid AN, Niazi AR. 2008. New records of graminicolous rust fungi from Pakistan.
Pak. J. Bot. 40(3): 1279-1283.
Ahmad §, Iqbal SH, Khalid AN. 1997. Fungi of Pakistan. Nabiza Printing Press, Karachi.
Ali A. 2006. Large scale production of sugarcane (Saccharum officinarum) through micropropagation
and in vitro selection of mutants resistant to Red rot disease in Sugarcane using induced
mutation. Ph.D. thesis, Department of Botany, University of the Punjab, Pakistan.
Butler EJ. 1914. Notes on some rusts in India. Ann. Mycol. 12: 76-82.
Cope TA. 1982. Flora of Pakistan, Poaceae. (Ed. Nasir E, Ali SI). Shamim Printing Press, Karachi.
Cummins GB. 1971. The Rust Fungi of Cereals, Grasses and Bamboos. Springer Verlag, Berlin.
Heidelberg. New York.
Hiratsuka N, Sato S, Katsuya K, Kakishima M, Hiratsuka Y, Kaneko S, Ono Y, Sato T, Harada Y,
Hiratsuka T, Nakayama K. 1992. The Rust Flora of Japan. Tsukuba Shuppankai, Tsukuba.
Ito S. 1909. On the Uredineae parasitic on the Japanese Gramineae. J. Sapporo Agric. Coll. Japan
3: 244-245,
Ryan CC, Egan BT. 1989. Rust, pp. 189-210. In Diseases of Sugarcane, Major Diseases (Ed. Ricand
C, Egan BT, Gillaspie JAG, Hughes CG). Elsevier, Amsterdam.
Sydow H, Sydow P, Butler EJ. 1906a. Fungi Indiae orientalis I. Ann. Mycol. 4: 422-445.
Virtudazo EV, Nojima H, Kakishima M. 2001. Taxonomy of Puccinia species causing rust diseases
on sugarcane. Mycoscience 42: 167-175.
:
:
MYCOTAXON
Volume 106, pp. 227-232 October-December 2008
Neotypification of Marasmius amazonicus
J.J.S. OLIVEIRA ', C. PUCCINELLI 7, M. CAPELARI ” & I.G. BASEIA !
‘jadson_oliver@yahoo.com.br — baseia@cb.ufrn.br
Depto. Botanica, Ecologia e Zoologia
Universidade Federal do Rio Grande do Norte
Campus Universitario, CEP: 59072-970, Natal, RN, Brazil
? puccinellic@yahoo.com.br — mcapelariibot@yahoo.com
Instituto de Botanica, Secdo de Micologia e Liquenologia
Caixa Postal 4005, CEP: 01061-970, Sao Paulo, SP. Brazil
Abstract — The holotype of Marasmius amazonicus was lost during the Second
World War due to the destruction of the Berlin-Dahlem herbarium, and, given the
nonexistence of isotypes, we propose a neotypification for the species.
Key words — Marasmiaceae, diversity, neotropics, taxonomy
Introduction
| The genus Marasmius was established by Fries’ Floram Scanicam, published
| in 1835-37, and currently contains around 2,000 widely distributed species.
| Marasmius amazonicus is one of the most beautiful members of the genus and
| belongs to series Leonini subsection Siccini section Sicci (Singer 1976). It was
_ originally collected near Jurua Miry, Amazonas State, Brazil, by Ule in 1901 and
sent to Germany, where Hennings (1904) described it as a new taxon, naming
| it after its native region. During the Second World War, the type specimen was
| lost after the destruction of the Berlin-Dahlem herbarium, where the original
| material was stored. Singer (1976) in his monograph of the tribe Marasmieae
_ for Flora Neotropica gave a detailed description of the species after material
| collected in Bolivia, where he commented on the loss of the type, although
| without designing a neotype.
This taxon was rediscovered in Atlantic Forest areas in Rio Grande do Norte
State, more precisely in Parque Estadual Dunas de Natal (Dunes State Park of
| Natal), a conservation area noted for its rich species diversity and interesting
_ discoveries (Baseia & Calonge 2005, Fazolino et al. 2008). Therefore, the
purpose of this paper is to establish a Brazilian neotype, providing a detailed
description with comments and illustrations of the species.
228 ... Oliveira & al.
Material and methods
The studied specimens were collected in the “Parque Estadual Dunas de Natal’,
a coastal Atlantic Forest reserve located in Natal city, Rio Grande do Norte
State, Brazil. The park covers an area of 1,172 ha. Average annual temperature
is around 26°C, and rainfall, between 800 mm and 1 555 mm per year, falling
predominantly between March and August. The fresh specimens were color-
coded according to Kornerup & Wanscher (1978) and described following the
usual techniques used in taxonomic studies of Marasmius (Gilliam 1976, Singer
1976, 1986; Desjardin & Petersen 1989, Desjardin et al. 2000, Wright & Albert6
2002, Wannathes et al. 2004).
Taxonomy
Marasmius amazonicus Henn., Hedwigia 43: 183. 1904.
PiLEus 16-58 mm diam., hemispheric to more often campanulate, central disc
smooth, slightly umbonate; margin decurved, sulcate; glabrous, but with a.
velutinous surface aspect, dry, membranous, hygrophanous, edge regular, purple
or deep lilac or dark “Magenta” (KW 13E8 13F8), with darker purple central
disc (KW 14F8) and presenting numerous buff dots which are oval or irregularly
rounded spots varying in diameter (2-3 mm), brownish yellow (KW 4A4) in
center and pale yellow (KW 4A3) at the margin. LAMELLAE free, sub-distant
(18-22) with no lamellulae, wide (2 mm broad), equal, neither intervenose nor
branched, pale yellow (KW 4A3), margin with regular edges and color similar
to pilear surface (KW 12F8), the surface inter lamellae similar in color to the
pileus. StrpE 50-70 x 1-2 mm, glabrous, smooth, shining, woody when dry
but resistant cartilaginous in young basidiomata, hollow, equal, central, terete
but sometimes compressed, blackish brown to chestnut color at the base and
pale cream at the apex, similar to lamellae color, though (KW 6F6-5E7), with
a well developed basal mycelium cream coloured (KW 3A2). CoNnTExT thin,
dextrinoid. BAstip1ospores 13.75-16.25(-17.5) x 2.5-3.75 um (Qm = 4.3, n
= 20 basidiospores), fusoid, smooth, hyaline, thin-walled, inamyloid (Fig. 2c).
BASIDIA not noticed. PLEUROCYSTIDIA absent. CHEILOCYSTIDIA similar to the
broom cells of the pilear surface, main body cylindric-clavate, hyaline, thin
to slightly thick-walled, 10-15 x 5-6.25 um; apical setulae 5-7.25 um broad
(Fig. 2b). HYMENOPHORAL TRAMA regular, dextrinoid, with hyaline hyphae,
thin-walled, septate, with small clamp connections but not noticed at all septa,
5-7.5 um diam. PILEIPELLIS hymeniform, composed by broom cells of the
Siccus-type, cylindrical to clavate, main body, hyaline when individualized and
slightly chestnut color when in group, thick-walled, 10-16.25 x 3.75-7.5 um,
apical setulae 3.75-5 um broad (Fig. 2a). Habitat: growing on decaying leaves.
Marasmius amazonicus neotypified ... 229
basidiomata (from neotype).
?
Fic. 1. Marasmius amazonicus
230 ... Oliveira & al.
Fra. 2. Marasmius amazonicus (from neotype).
a. broom cells. b. Cheilocystidia. c. Basidiospores.
MATERIAL EXAMINED: BRAZIL, RN, Natal: Parque Estadual Dunas de Natal, 27.V.2006,
JS. Oliveira, I.G. Baseia, B.D.B. Silva (Neotypus hic designatus UFRN-25 1); 11.V1.2007,
J.J.S. Oliveira, I.G. Baseia s.n. (UFRN); 14.V1.2007, J.J.S. Oliveira, I.G. Baseia s.n.
(UFRN).
Marasmius amazonicus neotypified ... 231
REMARKS—Marasmius amazonicus is a very peculiar species of Marasmius. It
is very well characterized by the bright coloration, predominantly purple with
white and cream-colored dots covering the entire pileus surface (Fig. 1). When
compared to a description by Singer (1976), the specimens studied here differ
in the umbonate pileus shape, corroborating Henning’s original description, in
contrast to the Bolivian species, described by Singer as umbilicate. Color photos
of this species can be found at the website: http://www.cb.ufrn.br/atlasvirtual/
Marasmius%20amazonicus.htm.
The neotype was collected in Atlantic Forest remnants, though several
studies proved its connectivity with Amazon Rainforest during the Tertiary
and Quaternary periods, a fact that could explain their high similarity (Prance
1987). Despite some expeditions to collect M. amazonicus in its original
location, Amazonian Rainforest, it was not possible to get any exemplar. The
main reason for choosing these Brazilian specimens instead of those from
the Singer collection, originating in Bolivia, was the advantage of being able
to describe the neotype from fresh, higher quality specimens that offered
better visualization of various important taxonomic features of the group,
such as the color pattern of the pileus, the basal mycelium and the most fragile
microstructures, which are more easily visualized in recent material. Thus, we
propose to establish the neotype, collected in the Parque Estadual Dunas de
Natal, in the state of Rio Grande do Norte, on the coast of Northeast Brazil.
|
: Acknowledgments
This research is supported by the Conselho Nacional de Desenvolvimento Cientifico e
Tecnoldgico (CNPq), Brazilian agency. The authors thank Professor Dennis E. Desjardin
_and Vladimir Antonin, for their useful comments, and Tereza Cristina Galvao for the
| drawing.
Literature cited
| Baseia IG, Calonge FD. 2005. Aseroé floriformis, a new phalloid with a sunflower-shaped receptacle.
Mycotaxon 92: 169-172.
| Desjardin DE, Petersen RH. 1989. Studies on Marasmius from eastern North America. 2. New
species. Mycotaxon 34(1): 71-92.
| Desjardin DE, Retnowati A, Horak E. 2000. Agaricales of Idonesia. 2. A preliminary monograph of
Marasmius from Java and Bali. Sydowia 52(2): 92-193.
Fazolino EP, Calonge FD, Baseia IG. 2008. Geastrum entomophilum, a new earthstar with an
unusual spore dispersal strategy. Mycotaxon 105 (in press).
Gilliam MS. 1976. The genus Marasmius in the Northeastern United States and adjacent Canada.
Mycotaxon 4: 1-144.
Hennings P. 1904. Fungi amazonici I. a cl. Ule collecti. Hedwigia 43: 154-186.
| Kornerup A, Wanscher JH. 1978. Methuen handbook of colours. Third edition. Eyre Methuen.
London. 252 pp.
232 ... Oliveira & al.
Prance GT. 1987. Biogeography of Neotropical plants. Biogeography and Quaternary history in
tropical America (ed. TC Whitmore, GT Prance), Clarendon Press, Oxford, pp. 46-65.
Singer R. 1976. Marasmieae (Basidiomycetes — Tricholomataceae). Flora Neotropica 17: 1-347.
Singer R. 1986. The Agaricales in Modern Taxonomy. 4th ed. Koenigstein: Koeltz Scientific Books,
981 pp.
Wannathes N, Desjardin DE, Retnowati A, Tan YS, Lumyong S. 2004. A redescription of Marasmius
pellucidus, a species widespread in South Asia. Fungal Diversity 17: 203-218.
Wright JE, Alberté E. 2002. Guia de los Hongos de la Regién Pampeana. I. Hongos con laminillas.
Buenos Aires: Literatura of Latin America, 279 pp.
MYCOTAXON
Volume 106, pp. 233-236 October-December 2008
Ochrolechia pallentiisidiata, a new species from China
ZE-FENG JIA**, QIANG REN* & ZUN-TIAN ZHAO”
zffia2008@163.com rengiang@sdnu.edu.cn ztzhao@sdnu.edu.cn
' College of Life Sciences, Shandong Agricultural University
Taian 271018, China
* College of Life Sciences, Shandong Normal University, Jinan 250014, China
Abstract — A new species of the lichen genus Ochrolechia, O. pallentiisidiata is
described from China. It is characterized by knobby-verrucose isidiata, having heavily
white pruinose apothecia, large ascospores and only apothecial margin KC+ red. The
species is collected from Yunnan Province in China.
Key words — taxonomy, lichenized ascomycetes, Ochrolechiaceae
Introduction
| During a study of the lichen family Ochrolechiaceae (Pertusariales, Ascomycota)
_ in China, an epiphloedal species new to science was found. The species differs to
_ other isidiate species in having a larger ascospores and heavily white pruinose
_ apothecia.
Material and methods
| The specimens examined collected from China are preserved in the Lichen
| Section of Herbarium Mycologicum Instituti Microbiological Academine
| Sinicae (HMAS-L).
| For the morphological and anatomical observations were made using
| a dissecting microscope (TECK-XTS) and a compound microscope
(OLYMPUS-CHB) for all materials. Hand-sections were routinely made and
examined with water used as the mounting medium. Chemical substances
_ were studied by the color tests with 10% KOH (K), sodium hypochlorite (C)
and alcoholic solution of p-phenylenediamine (PD), and means of thin-layer
| chromatography (Culberson & Kristensson 1970, Culberson 1972).
# Ze-Feng JIA and Qiang REN contributed equally to this work
* Corresponding author
234 ... Jia, Ren & Zhao
Fic. 1 Ochrolechia pallentiisidiata.
A. Habit (bar = 1 mm); B. Two asci containing ascospores (Bar=100um)
Taxonomic description
Ochrolechia pallentiisidiata Z.F. Jia & Q. Ren, sp. nov. ~ FIGURE 1
MycoBank MB 512228
Similis Ochrolechia pallescens sed isidiosus, acidum gyrophoricum cotinenti et ascosporis
majoribus differt.
Typus: CHINA, Yunnan Provincia, Weixi, Yezhi, Baimaluo, alt. 2400m, Jing-Jun Su 0241,
8.v.1982; holotype: HMAS-L.
EtyMoLoGcy: ‘The specific epithet pallentiisidiata is derived from the Latin participle
pallens, meaning pale, referring to the white pruinose apothecial disks; and the adjective
isidiatus referring to the isidiate thallus.
THALLUS crustose, adnate, up to 1.0 mm thick, becoming thin at edge; grey
to dark grey, rough, verruculose to verrucose; knobby to verrucose isidia, the
isidium 0.2-0.4 mm across and up to 0.8 mm long, sometimes growing large,
concolorous with the thallus. APOTHECIA sessile, dispersed, roundish, dish-
shaped, 1-3 mm in diameter; discs pinkish to orange-pinkish when young,
then heavily white pruinose on surface when mature, slightly concave to plane;
apothecial margin entire, smooth to verrucose, but not isidiate, 250-400 um
thick, concolorous with the thallus.
APOTHECIAL ANATOMY: Hymenium hyaline, 300-350 um high; epithecium
brown, 20-40 um thick; hypothecium slightly brownish, 50-70 um; paraphyses
dense and netted, 1.5-2.0 um wide; excipulum proprium absent; amphithecium
moderately thick, well-developed medulla; cortex 30-50 um laterally, expanding
to over 150 um at base; algal layer usually forming a continuous layer below
hypothecium, scattered to margin. Asci long claviform, 6-8 spores per ascus;
ascospores hyaline, simple, ellipsoid to broadly ellipsoid, 70-85 x 37.5-45 um.
CHEMIsTRY: Thallus cortex and medulla K-, C-, KC-, PD-; apothecial disc
and margins K-, C+ red or C-, KC+ red, PD+; UV-; containing gyrophoric
acid (major) and lecanoric acid.
239
‘IAT]LSIU IO JUNSGe= — ‘OAT]ISOd JO JUdsaId= + :sjoquidsS
Ochrolechia pallentiisidiata sp. nov. (China) ...
= = = ie _ = = QUOYJUPXIYOTT
- _ + - - + = proe o11ejOLIeA,
= a fe is oe _ ae proe s110uR27T
+ + = + if + proe s10ydoiky
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- “ 4 a _ = - ey[Npowi
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NOILLOVaY 4)
GuG VE-OC 6I-F1 Goer Gll VE-ST O€-07 GV-SLE (wm)
x 1258S x 19-PP SALSA AG X VV-LT xX LTO-TE X €9-8P x 8-02 SaUuOdSOOSY
dsoutmnid esoutnid dsourini ssoutmnid esoutnid dsoutnid ait ssournid euIni
ae Apysry a oe Anysys ee Goyrgeiaeari ea
O1-S'T (0'€-)0°7-6'0 S7-S'T 06-0 | O'€-0'1 (0'b-)0°€-0'1 O'€-0'1 (vuu) “wei
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0} dAeDU07) 0} aAvou07) Id Id Id : Id et o x
Te[nuess 0} Jepnuri3s= Jepnueis Juaystsiad uaystsiad 7 ISOONIIIA baa
Aqqouy “apueys eIpIsiopnasg 0} Aqqouy ‘Iapugys ‘Aqqouy Ty], 0} Aqqouyy PIS]
sso Sno]oor10 snojool1o snojoo}40 eae ae snojoo40 snojoon40 SATIVH
W JOIA0Z) [OIA0D) [OIHAOD) 29 snojosT105 JOINAOD JOIHAOD LL
SISNAIDVNV WOUVTTILLNV V.LVIGISIAAS V.LVICISI AVGOSVA SNAOSATIVd V.LVICISILLNATTVd SHIOadS
sardods Iv[IUIs YIM vyvIpIsiUavd V1IYIIJOLYIO JO UOTsTIVdWIOD *[ ATAVI,
236 ... Jia, Ren & Zhao
HABITAT AND DISTRIBUTION: The species is so far known only from the type
locality where it grows on the bark at the base of Abies fabri (Mast.) Craib,
Yunnan Province, southern China.
Ochrolechia pallentiisidiata is characterized by having knobby to verrucose
isidia, large apothecia 1-3 mm in diam., disks covering white pruinose, asci
6-8-spored, large ascospores 70-85 x 37.5-45 um and mainly present
gyrophoric acid. It is similar to O. pallescens (L.) A. Massal. in thallus and
white pruinose apothecia, but differs in having isidia, larger ascospores and
chemistry (Verseghy 1962). The new species also differs from other isidiate
species in having larger ascospores and somewhat different isidia and chemistry
(Verseghy 1962, Howard 1970, Brodo 1991, Hanko et al. 1986). The diagnostic
features of the species and similar species are summarized in table 1.
ADDITIONAL MATERIAL EXAMINED: CHINA. Yunnan Province, Weixi, Weideng, Xinhua,
alt. 320m, Jing-Jun Su 0805, 24.v.1982.
Acknowledgments
The authors thank Prof A. Aptroot and X.G. Zhang for pre-submission reviews. This
project was supported by the National Natural Science Foundation of China (No.
30470003).
Literature cited
Brodo IM. 1991. Studies of the genus Ochrolechia. 2. Corticolous species of North America.
Canadian Journal of Botany 69: 733-772.
Culberson CE. 1972. Improved conditions and new data for the identification of lichen products by
a standardized thin-layer chromatographic method. Journal of Chromatography 72: 113-125.
Culberson CF, Kristensson H. 1970. A standardized method for the identification of lichen
products. Journal of Chromatography 46: 85-93.
Hanko B, Leuckert C, Ahti T. 1986. Beitrage zur Chemotaxonomie der Gattung Ochrolechia
(Lichenes) in Europa. Nova Hedwigia 42: 165-199.
Howard GE. 1970. The lichen genus Ochrolechia in North America north of Mexico. The Broyologist
73: 93=130.
Verseghy K. 1962. Die Gattung Ochrolechia. Beihefte zur Nova Hedwigia 1: 1-146.
MYCOTAXON
Volume 106, pp. 237-241 October-December 2008
Kretzschmaria albogrisea sp. nov.
and K. curvirima from Brazil
JADERGUDSON PEREIRA", JOSE Lurz BEZERRA' & LEONOR Costa MAIA?
*jader@uesc.br
Dept’. de Ciéncias Agrarias e Ambientais, Universidade Estadual de Santa Cruz - UESC
Rod. Ilhéus-Itabuna, km 16, Ilhéus, BA, 45662-000, Brazil
*Dept’. de Micologia, Universidade Federal de Pernambuco - UFPE
Av.Prof. Nelson Chaves s/n, Recife, PE, 50670-420, Brazil
Abstract — Kretzschmaria albogrisea is described as a new species. It is distinguished
primarily by its white and gray surface of the stroma, which has an unusual shape and
size. Occurrence of K. curvirima in Brazil is reported.
Key words — systematics, Xylariaceae, Ascomycota
Introduction
Kretzschmaria is a genus of Xylariaceae whose species are mainly found in
tropical and subtropical regions. Rogers & Ju (1998) accepted 16 taxa. Later, the
following taxa were included: K. orientalis Lar.N. Vassiljeva, K. argentinensis
Hladki & A.I. Romero, K. sigmoidirima Hladki & A.I. Romero and K. varians
J.D. Rogers & Y.M. Ju, (Vassiljeva 1998, Hladki & Romero 2001, Rogers & Ju
2004).
In Brazil, K. cetrarioides (Welw. & Curr.) Sacc., K. clavus (Fr.) Sacc., K. lucidula
(Mont.) Dennis, K. micropus (Fr.) Sacc., K. pavimentosa (Ces.) P.M.D. Martin,
K. sandvicensis (Reichardt) J.D. Rogers & Y.M. Ju and K. sigmoidirima have
already been reported (Theissen 1908, Sampaio 1916, Chardon et al. 1940,
Viégas 1945, Dennis 1957, Batista et al. 1966, Batista & Peres 1967, Silveira &
Rodrigues 1985, Rogers & Ju 1998, Hladki & Romero 2003, de Meijer 2006).
In this work we are proposing a new species of Kretzschmaria and reporting
the occurrence of K. curvirima in Brazil.
Materials and methods
On May 2008, a specimen of Kretzschmaria was collected in the “Reserva
Particular do Patriménio Natural Fazenda Santa Beatriz do Carnijo, an area
of Atlantic Rain Forest in the municipality of Moreno, State of Pernambuco,
Brazil. Asci and ascospores were observed in water and Melzer’s reagent and
238 ... Pereira, Bezerra & Maia
analyzed according to Rogers & Ju (1998). Attempts to culture this fungus
failed. The exsiccate was deposited in the mycological collection of the URM
Herbarium and the description registered in the MycoBank.
A specimen of K. curvirima from Amazonas, deposited in the URM
Herbarium, was also examined.
Taxonomy
Kretzschmaria albogrisea Jad. Pereira, J.L. Bezerra & L.C. Maia, sp. nov. Fics. 1-5
MycoBank MB512104
Stromata superficialia, solitaria vel gregaria, subglobosa vel hemisphaerica, usque ad 10
mm lata x 2.5-4.5 mm alta; extus alba et grisea, dura, levis; intus alba, mollia. Perithecia
subglobosa, 0.4-1 mm lata x 0.6-1 alta. Ostiola leviter elevata. Asci octospori, 245-312 um
longitudine tota x 10-11 um crassi, partibus sporiferis 175-217 um longitudine, annulo
apicali in liquore iodato Melzeri cyanescente, plus minusve cylindraceo, 10-11 ym alto x
5-5.5 um lato. Ascosporae fuscae, ellipsoideae vel oblongae, plus minusve inequilaterales,
levis, 31-37 x 9-13 um, rima germinativa recta longa vel brevi praeditae.
EryMo_oey: referring to the coloration of the stromatal surface.
Stromata superficial, solitary or gregarious, subglobose, depressed-spherical
to hemispherical, 3.5-10 mm diam x 2.5-4.5 mm high; tissue between and
beneath perithecia spongy, white; surface hard, white and gray with mottled
aspect. Perithecia spherical to obovoid, 0.4-1 mm diam x 0.6-1 mm high.
Ostioles papillate. Asci 8-spored, 245-312 um total length x 10-11 um broad,
spore-bearing part 175-217 um long, with the apical ring bluing in Melzer's
iodine reagent, urn-shaped, 10-11 um high, 5-5.5 um broad. Ascospores
uniseriate, blackish brown, ellipsoid, more or less inequilateral, smooth, 31-37
x 9-13 um, with straight germ slit spore-length to slightly less.
SPECIMEN EXAMINED: BRAZIL: Pernambuco, Moreno, Reserva Particular do Patriménio
Natural Fazenda Santa Beatriz do Carnijé (08°08'38.9"S and 35°04’35.3” W, 120m alt.),
13.V.2008, leg. Jad. Pereira, in bark of indeterminated tree (HOLOTYPE, URM 79233).
Notes: Kretzschmaria albogrisea differs from all species of the genus by the
sessile stroma with white and gray surface. The species K. micropus has similar
straight germ slit and ascospore size, but presented blackish brown stromata
covered with spines and conical ostioles (Rogers & Ju 1998). Although
K. macrosperma has straight germ slits and a similar format of ascospores, it
differs from K. albogrisea by its large ascospores (45-67 x 12.5-17.5 um) and
different stroma.
Kretzschmaria curvirima J.D. Rogers & Y.M. Ju, Mycotaxon 68: 355. 1998.
FIGs. 6-8
Stromata carbonaceous with convex, obconical fertile parts 4.5-6.5 x 4-5.2
mm diam, 1.9-4.3 mm high, containing one or two perithecia, aggregated or
Kretzschmaria albogrisea sp. nov. (Brazil) ... 239
Fics. 1-5: Kretzschmaria albogrisea. 1. Stromata. 2. Perithecia. 3. Ascospores released from the
Ostioles (arrow). 4. Ascospore with straight germ slit slightly smaller than spore-length. 5. Ascus
apical ring colored in Melzer’s reagent. 6-8: Kretzschmaria curvirima. 6. Stroma. 7. Ascospores
with sigmoid germ slit. 8. Ascus apical ring colored in Melzer’s reagent (arrow).
Seale bars)’ =44mm;)2'= 2 mm;73 = 200 fim’; 6"= 3mm: 4,577) 3 = 10 um:
solitary, bearing one to several conic spines on top; surface dull blackish brown;
tissue between and beneath perithecia coriaceous, white to gray, becoming dark
brown and disintegrating. Perithecia obovoid, 0.5-2 mm diam x 0.9-1.7 mm
240 ... Pereira, Bezerra & Maia
high. Ostioles conical. Asci damaged, with the apical ring bluing in Melzer’s
reagent, urn-shaped, 10-10.5 tm high, 5-5.5 um broad. Ascospores dark
brown, unicellular, ellipsoid-inequilateral, with narrowly to broadly rounded
ends sometimes pinched, smooth, (35.5—)36-50 x (8—)8.5-16 um, with sigmoid
to spiral germ slit slightly smaller than spore-length.
SPECIMEN EXAMINED: BRAZIL: Amazonas, Manaus, Reserva Ducke, 25.1II.1961,
leg. G. E. P. Peres, on wood (URM 21237, as Kretzschmaria spinifera).
Notes: Rogers & Ju (1998) and Hladki & Romero (2001) reported, respectively,
K. curvirima and K. sigmoidirima with sigmoid and spiral ascospore germ
slit. The specimen examined in this work presented similar characters to
K. curvirima, differing from K. sigmoidirima by the presence of conic spines
in the stromata and larger ascospores. Batista and co-workers identified this
specimen as K. spinifera but did not report in their papers. Viégas (1945)
reported a specimen correctly identified by Julian H. Miller as K. spinifera (=
K. micropus) in the State of Sao Paulo.
Acknowledgments
Thanks are due to Dr. Jack D. Rogers and Dr. Adriana I. Hladki for reviewing the
manuscript and making valuable suggestions. The authors are indebted to the Fundac¢ao
de Amparo a Pesquisa do Estado da Bahia (FAPESB) and to the Conselho Nacional de
Desenvolvimento Cientifico e Tecnol6gico (CNPq) for financial support. Thanks are
also due to RPPN Fazenda Santa Beatriz do Carnij6 manager for allowing the field trips
and Dr. Tatiana B. Gibertoni for advice during preparation of the manuscript.
References
Batista AC, Falcao RGS, Peres GEP, Moura NR. 1966. Fungi Paraenses. Revisao da cole¢ao de
Paul Hennings, do Museu Paraense Emilio Goeldi. Publicacées. Instituto de Micologia da
Universidade do Recife 506: 1-290.
Batista AC, Peres GEP. 1967. Xylosphaeraceae: estudo de algumas espécies. Atas do Instituto de
Micologia. Universidade Federal de Pernambuco, Recife 5: 115-130.
Chardon CE, Miller JH, Muller AS. 1940. Ascomycetes from the State of Minas Gerais (Brazil).
Mycologia 32: 172-204.
Dennis RWG. 1957. Further notes on tropical American Xylariaceae. Kew Bull. 2: 297-332.
Hladki Al, Romero AI. 2001. The genus Kretzschmaria from Tucuman, Argentina. Mycotaxon 79:
481-496.
Hladki AI, Romero AI. 2003. Two new species of Stilbohypoxylon and the taxonomic positions of
Hypoxylon cyclopicum, H. chionostomum and Anthostoma chionostoma. Sydowia 55: 65-76.
Meijer AAR de. 2006. Preliminary list of the macromycetes from the Brazilian State of Parana. Bol.
Mus. Bot. Mun. 68: 1-55.
Rogers JD, Ju Y-M. 1998. The genus Kretzschmaria. Mycotaxon 68: 345-393.
Rogers JD, Ju Y-M. 2004. Kretzschmaria varians sp. nov., Xylaria coremiifera sp. nov. and Xylaria
umbonata sp. nov. from Costa Rica. Mycol. Progress 3: 37-40.
Sampaio AJ. 1916. A flora de Mato Grosso. Arch. Mus. Nac. 19: 1-126.
Kretzschmaria albogrisea sp. nov. (Brazil) ... 241
Silveira VD, Rodrigues KF. 1985. Levantamento preliminar de Xylariaceae da Amazénia. Acta
Amazonica 15: 7-27.
Theissen F. 1908. Fragmenta brasilica I. Annls. mycol. 6: 531-535.
Vassiljeva LN. 1998. Nizshie Rasteniya, Griby i Mokhoobraznye Dalnego Vostoka Rossii, Griby.
Tom 4. Pirenomitsety i Lokuloaskomitsety. (Sankt-Peterburg): 195.
Viégas AP. 1945. Uns poucos fungos do Brasil. Bragantia 5: 561-570.
MYCOTAXON
Volume 106, pp. 243-246 October-December 2008
Biogeography and hosts of poroid wood decay fungi in
North Carolina: species of Trametes and Trichaptum
L.E GRAND, C.S. VERNIA & M.J. MUNSTER
larry_grand@ncsu.edu
Department of Plant Pathology, North Carolina State University
Raleigh, North Carolina 27695-7616 USA
Abstract— Distribution and host plants are provided for seven species of Trametes and
six species of Trichaptum. Twenty three of the host-fungus combinations reported here
are new records for the United States. County distribution maps are provided for each
species as well. The complete checklist can be found at:
www.cals.ncsu.edu/plantpath/people/faculty/grand/projects/mycotaxon_6.pdf
Keywords—fungus distribution, polypores
Introduction
Previous studies by Grand & Vernia (2004a,b) addressed the importance of
biodiversity and biogeography of fungi in ecosystems of North Carolina. The
occurrence of selected genera and host plants of poroid wood decay fungi in
North Carolina have been reported by Jung (1987), Vernia & Grand (2000)
and Grand & Vernia (2002, 2005a,b, 2007). These studies have expanded the
geographical range and host plant associations of poroid wood decay fungi in
the southern region of the United States. This report is the sixth in a series in a
long-term study of poroid wood decay fungi in North Carolina.
Materials and methods
Poroid wood decay fungi were collected in North Carolina from 1997-2007.
Collections were made of all species of Trametes and Trichaptum on new or
unusual hosts. Specimens were placed in paper bags in the field with a sample of
decayed wood for the majority of collections and field notes for all collections.
Specimens were examined in the laboratory and identified using the taxonomic
treatments of Gilbertson & Ryvarden (1987), Jung (1987) and Overholts (1953).
Nomenclature and authorities are from Index Fungorum (CABI et al. 2007) for
fungi and Kartesz (1994) and IPNI (2007) for host plants.
244 ... Grand, Vernia & Munster
The majority of collection sites were in state parks, game lands and natural
areas, the Nantahala, Pisgah, Croatan and Uwharrie National Forests and the
Blue Ridge Parkway and Great Smoky Mountains National Parks. A county
distribution map is provided for each species (Figs. 1-12).
Data from previous studies (Jung 1987, Grand et al. 1975), collections in
the mycological herbarium (NCSC) at North Carolina State University (NCSU)
and the University of Tennessee Mycological Herbarium (Ma & Petersen 2007),
records of the Plant Disease and Insect Clinic, Plant Pathology Department,
NCSU, and the BPI web site (Farr et al. n.d.) were used in developing distribution
maps.
Results and Discussion
Seven species of Trametes were recorded. Trametes elegans (Spreng.) Fr. (Fic. 3),
T: hirsuta (Wulfen) Pilat (Frc. 4), and T. versicolor (L.) Lloyd (Fic. 6) are common
and widespread throughout North Carolina. The limited distribution of T.
conchifera (Schwein.) Pilat (Fic. 2) does not reflect the statewide distribution of
its most common host, elm (Ulmus spp.). One would expect T. conchifera to be.
more widespread throughout the state. Trametes pubescens (Schumach.) Pilat
(Fic. 5) was recorded from 13 counties in all three physiographic provinces.
Trametes cervina (Schwein.) Bres. (Fic. 1) and T: villosa (Sw.) Kreisel (Fic. 7)
were not collected frequently enough to determine a distribution pattern.
Six species of Trichaptum were recorded. Trichaptum abietinum (Pers.)
Ryvarden (Fic. 8) and T! biforme (Fr.) Ryvarden (Fic. 9) are widespread in North
Carolina. Trichaptum abietinum occurs only on conifers, and T: biforme occurs
almost exclusively on deciduous trees. Trichaptum fuscoviolaceum (Ehrenb.)
Ryvarden (Fic. 11) was found in nine counties in all three physiographic
provinces. Trichaptum byssogenum (Jungh.) Ryvarden (Fic. 10) and T
laricinum (P. Karst.) Ryvarden (Fic. 12) were not collected frequently enough
to determine a distribution pattern. Trichaptum sector (Ehrenb.) Kreisel (Fic.
12) was found in nine counties in the Coastal Plain. It is typically a southeastern
species (Gilberston & Ryvarden 1987) and may reach the northern limits of its
range in North Carolina.
Fic. 1. Distribution of Trametes cervina Fic. 2. Distribution of Trametes conchifera
in North Carolina. in North Carolina.
Trametes & Trichaptum in North Carolina ... 245
Fic. 3. Distribution of Trametes elegans Fic. 4. Distribution of Trametes hirsuta
in North Carolina. in North Carolina.
Fic. 5. Distribution of Trametes pubescens Fia. 6. Distribution of Trametes versicolor
in North Carolina. in North Carolina.
Nd
Fic. 7. Distribution of Trametes villosa Fic. 8. Distribution of Trichaptum abietinum
in North Carolina. in North Carolina.
Fic. 9. Distribution of Trichaptum biforme Fic. 10. Distribution of Trichaptum
in North Carolina. byssogenum in North Carolina.
Fic. 11. Distribution of Trichaptum Fic. 12. Distribution of (A) Trichaptum
fuscoviolaceum in North Carolina. laricinum and (B) T. sector in North Carolina.
246 ... Grand, Vernia & Munster
Acknowledgements
The authors thank Drs. Richard Baird and Lauraine Hawkins for their suggestions and
comments that improved the manuscript. A special thanks to Tom Howard and the
staff of the North Carolina State Parks system for permission to collect in the parks
and natural areas. The loan of collections from the University of Tennessee Herbarium
(TENN) is gratefully acknowledged. Financial support for this project was provided
in part by generous grants from the Highlands Biological Station, Highlands, North
Carolina.
Literature cited
CABI Bioscience, Centraalbureau voor Schimmelcultures, Landcare Research (custodians). 2007.
Index Fungorum. Published on the Internet http://www.indexfungorum.org [accessed 14 & 18
June 2007].
Farr DE, Rossman AY, Palm ME, McCray EB. (n.d.) Fungal Databases, Systematic Botany
& Mycology Laboratory, ARS, USDA. Published on the Internet http://nt.ars-grin.gov/
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MYCOTAXON
Volume 106, pp. 247-267 October-December 2008
Glomus irregulare, a new arbuscular mycorrhizal fungus
in the Glomeromycota
JANUSZ BEASZKOWSKI, BEATA CZERNIAWSKA
janusz. blaszkowski@agro.ar.szczecin.pl
Department of Plant Protection, University of Agriculture
Stowackiego 17, 71434 Szczecin, Poland
TESFAYE WUBET, TINA SCHAFER, FRANCOIS BUSCOT
tesfaye. wubet@ufz.de
Helmholtz Centre for Environmental Research - UFZ
Theodor-Lieser-Strafse 4, 06120 Halle-Saale, Germany
CARSTEN RENKER
Carsten. Renker@stadt.mainz.de
Naturhistorisches Museum Mainz, Landessammlung fiir Naturkunde Rheinland-Pfalz
Reichklarastr. 10, D-55116 Mainz, Germany
Abstract — A new arbuscular mycorrhizal fungus (Glomeromycota) is described and
illustrated. Glomus irregulare was found associated with different plants colonizing
maritime sand dunes of Giftung Island, Egypt (Africa), Bornholm (Denmark), Greece,
Poland, and Spain. This fungus produces spores aggregated inside roots or in the soil
or (rarely) singly in the soil. Spores form blastically either (when aggregated) at the tip
of or along dichotomously branched hyphae continuous with mycorrhizal extraradical
hyphae or (when single) terminally from the unbranched extraradical hyphae. The
hyaline to pale yellow spores are ovoid, oblong to irregular, 60-130 x 80-240 um, and
may have deep wall depressions and apical cap-like swellings. The spore wall consists
of two semi-permanent, hyaline outer layers and a hyaline to pale yellow, laminate
innermost layer staining in Melzer’s reagent. Glomus irregulare formed vesicular-
arbuscular mycorrhizae in single-species cultures with Plantago lanceolata as the host
plant. Phylogenetic analyses do not yet conclusively support the obvious morphological
and biochemical differences between G. irregulare and G. intraradices, and the need to
sequence other rDNA regions to further refine genetic relationships is suggested.
Key words — distribution, molecular phylogeny
Introduction
Arbuscular mycorrhizal fungi (AMF), which are believed to have evolved
over 450 million years ago, associate with most vascular land plants and have
248 ... Blaszkowski & al.
a worldwide distribution (Redecker et al. 2000, Smith & Read 1997). Among
the many beneficial effects of the symbiosis between AMF and plants are the
increased productivity and strength of mycorrhizal plants and the positive
influence of these fungi on plant biodiversity (Smith & Read 1997, van der
Heijden et al. 1998).
For many years, AMF were placed in phylum Zygomycota, first in the family
Endogonaceae Paol. of the order Endogonales Moreau (Gerdemann & Trappe
1974, Morton & Benny 1990a), and then in the order Glomerales J.B. Morton
& Benny (Morton & Benny 1990a). However, molecular analyses have revealed
that these fungi have no affinity with either members of the Zygomycota or
other known major fungal groups (SchiiBler et al. 2001). Consequently, they
were removed from the Zygomycota to be accommodated in a new phylum, the
Glomeromycota C. Walker & A. Schiissler, presently comprising four orders,
ten families, and fourteen genera (Blaszkowski 2003, Oehl & Sieverding 2004,
Palenzuela et al. 2008, SchiiBler et al. 2001, Sieverding & Oehl 2006, Walker &
A. Schissler 2004, Walker et al. 2007a, b, 2008). |
The most numerous and diverse fungal group in the Glomeromycota is
the genus Glomus Tul. & C. Tul., comprising ca. 53% of all arbuscular fungi
described to date. Within Glomus, Schwarzott et al. (2001) established three
phylogenetic groups, Glomus groups A, B, and C. Walker & SchtiBler (2004)
elevated Glomus group C to the rank of genus, Diversispora C. Walker & A.
Schiissler, in the newly erected family Diversisporaceae C. Walker & A. Schiissler
and order Diversisporales C. Walker & A. Schiissler. The Diversisporales now
includes four additional families, the Acaulosporaceae J.B. Morton & Benny,
Entrophosporaceae Oehl & Sieverd., Gigasporaceae J.B. Morton & Benny, and
Pacisporaceae C. Walker et al. (Blaszkowski 2003, SchiiBler & Walker 2004,
Sieverding & Oehl 2006). Whereas Glomus group B was phylogenetically
uniform, Glomus group A contained two or three subclades (a-c), with subclade
“c” including only one undescribed Glomus sp. isolate (W3347; Schwarzott et
al. 2001) to which many other sequences have been added (Kovacs et al. 2007).
Representatives of the other subclades include G. mosseae (T.H. Nicolson &
Gerd.) Gerd. & Trappe (subclade “a’) and G. intraradices N.C. Schenck & GS.
Sm. (subclade “b”)
Apart from interspecific genetic differences, AMF also exhibit intraspecific
divergences, especially within Glomeraceae Piroz. & Dalpe. Antoniolli et al.
(2000) found a 3-5-fold higher genetic diversity within G. mosseae spores
than in Gigaspora margarita W.N. Becker & IR. Hall that was isolated from
the same site. Jansa et al. (2002), who analyzed intra- and intersporal diversity
using the internal transcribed spacer (ITS) of ribosomal DNA sequences from
three G. intraradices isolates, revealed two different sequence types within a
single-spore isolate. Moreover, phylogenetic analyses of all sequences known
Glomus irregulare sp. nov. ... 249
at that time, including those deposited in GenBank, showed four distinct ITS
sequence types for G. intraradices.
Investigations of AMF occurrence in rhizosphere soil and root samples
collected from different regions in Europe revealed a frequent irregularly-
shaped Glomus spore with a morphology unlike that found in any other
described Glomus species. Although this fungus tended to form intraradical
spores similar to G. intraradices and its extraradical globose spores superficially
resembled those from lighter-coloured isolates of the latter fungus, marked
differences mainly in (1) the mode of spore formation, (2) aggregated spore
organization, and (3) the phenotypic and biochemical properties of their spore
wall components convinced us that these fungi are morphologically distinct.
Therefore, the newly discovered Glomus morphotype is described below as G.
irregulare sp. nov. Phylogenetic analyses of sequences from G. irregulare spores,
however, indicate that they partly cluster with G. intraradices sequences. The
contradiction between the morphological and molecular analyses of Glomus
irregulare and G. intraradices is discussed in detail below.
Materials and methods
Establishment and growth of trap and single-species cultures, extraction of
spores, and staining of mycorrhizae
Spores examined in this study came from both pot trap and single-species cultures.
Trap cultures were established to obtain a great number of living spores and to initiate
sporulation of species that were present but not sporulating in the field collections (Stutz
& Morton 1996). The method used to establish trap cultures, their growing conditions,
and the methods of spore extraction and staining of mycorrhizae were as those described
previously (Blaszkowski et al. 2006).
Single-species cultures were also generally established and grown as given in
Blaszkowski et al. (2006), with two exceptions. First, instead of marine sand, their
growing medium was an autoclaved commercially available coarse-grained sand (grains
1.0-10.0 mm diam — 80.50%; grains 0.1-1.0 mm diam - 17.28%; grains < 0.1 mm diam
— 2.22%) mixed (5:1, v/v) with clinopthilolite (Zeocem, Bystré, Slovakia) of grains 2.5-5
mm. Clinopthilolite is a crystalline hydrated alumosilicate of alkali metals and alkaline
earth metals having, e.g., a high ion exchange capability and selectivity, as well as a
reversible hydration and dehydration. pH of the sand-clinopthilolite mixture was 7.3.
Second, the cultures were kept in transparent plastic bags, 15 cm wide and 22 cm high as
suggested by Walker & Vestberg (1994), rather than open pot cultures (Gilmore 1968).
To prevent contamination of the cultures with other AMF but still to allow exchange
of gases, we left an opening, about 1 cm wide, in the centre of the upper part of each
bag, while the edges on both sides were closed with small plastic clips. The cultures
were watered with tap water once a weak, harvested after five months when spores were
extracted for study. To reveal mycorrhizae, root fragments located ca. 1-5 cm below the
upper level of the growing medium were cut off with a scalpel. Plantago lanceolata was
the host plant used in both trap and single-species cultures.
250 ... Blaszkowski & al.
Microscopy survey
Morphological properties of spores and their wall structure were determined based
on examination of at least 100 spores mounted in polyvinyl alcohol/lactic acid/glycerol
(PVLG; Omar et al. 1979) and a mixture of PVLG and Melzer's reagent (1:1, v/v). Spores
at all developmental stages were crushed to varying degrees by applying pressure to the
cover slip and then stored at 65°C for 24 h to clear their contents from oil droplets. They
were then examined under an Olympus BX 50 compound microscope equipped with
Nomarski differential interference contrast optics. Microphotographs were recorded on
a Sony 3CDD color video camera coupled to the microscope. |
Terminology of spore structure is that suggested by Stiirmer & Morton (1997) and
Walker (1983). Spore colour was examined under a dissecting microscope on fresh
specimens immersed in water. Colour names are from Kornerup & Wanscher (1983).
Nomenclature of fungi and plants is that of Walker & Trappe (1993) and Mirek et al.
(1995), respectively. The authors of the fungal names are those presented at the Index
Fungorum website _http://www.indexfungorum.org/AuthorsOfFungalNames.htm.
Voucher specimens were mounted in PVLG and a mixture of PVLG and Melzer’s reagent
(1:1, v/v) on slides and deposited in the Department of Plant Protection, University of
Agriculture, Szczecin, Poland, and in the herbarium at Oregon State University (OSC) |
in Corvallis, Oregon, U.S.A.
Colour microphotographs of spores of G. irregulare can be viewed at the URL http://
www.agro.ar.szczecin.pl/~jblaszkowski/.
Spore cleaning and DNA extraction
Spores of G. intraradices, G. irregulare, and G. geosporum (the last species selected
to represent an outgroup in phylogenetic analyses) used in the molecular analyses were
isolated from single-species cultures established from spores produced in trap cultures
with mixtures of the rhizosphere soil and root fragments of Ammophila arenaria (L.)
Link growing in maritime dunes of Bornholm, Denmark. Spores were separated in a
Petri dish with fresh and autoclaved double distilled water using a pipette. Spores (ca.
10) were transferred to 1.5 ml reaction tubes and cleaned by washing in sterile filtered
SDS (1%) and rinsing twice with fresh double distilled water followed by washing in
absolute ethanol. The tubes were air dried under the clean bench and kept at -20°C
until DNA extraction. Spore DNA was extracted following the procedure described by
Manian et al. (2001). The samples were frozen in liquid nitrogen in 1.5 ml tubes and
crushed thoroughly using a micropestle. To each sample, 100 ul of lysis buffer (200 mM
TrisHCl pH 7.5, 250 mM NaCl, 1 mM EDTA and 1% SDS, w/v) was added and mixed
thoroughly. The samples were subjected to three alternate cycles of freezing in liquid
nitrogen and incubation at 100°C for 1 min, followed by freezing and a final incubation
at 100°C for 10 min. The contents of each tube were centrifuged at 11 000 g for 3 min
and the supernatant collected carefully. The supernatants were passed through DNA
purification columns (QIAquick, Qiagen) according to the manufacturer's instructions.
The purified DNA was eluted in 50 ul of 1 mM TrisHCl pH 8.5 prewarmed to 55°C.
PCR amplification
The nuclear ribosomal SSU and ITS rDNA genes were amplified with two parallel
nested PCR reactions. Partial SSU rDNA was amplified using the primer pair
Glomus irregulare sp. nov. ... 251
GlomerWT0 (5’-CGA TGD WTC ATT CAA ATT TCT GCC -3’) and Glomer1536 (5’-
RTT GCA ATG CTC TAT CCC CA-3’) in the first PCR amplification and GlomerW T0
and GlomerWT2 (5’°-CAA ACT TCC ATC GGT TAR ACA CCG-3’) for the second
PCR (Wubet et al. 2006). The end part of the SSU with the complete internal transcribed
spacer (ITS) region was amplified using the primer set NS5 (5° -AAC TTA AAG GAA
TTG ACG GAA G- 3’) and ITS4 (5° -TCC TCC GCT TAT TGA TAT GC- 3’) for the
first PCR and Glom1310 (5° -AGC TAG GYC TAA CTT TGT TA- 3’) and ITS4 for the
nested PCR amplification (Redecker 2000). In both steps, DNA was amplified in 50 ul
reaction volume containing PCR buffer (Invitrogen, Karlsruhe), concentrations of 0.2
mM of each dNTP (Invitrogen) and 0.5 mM of each primer, 1.5 units of recombinant
Taq DNA polymerase (Invitrogen), and 1 ul of DNA template, using the program Touch
60. A 4 ul aliquot of each nested PCR product was analysed on a 1% agarose gel followed
by ethidium bromide (0.5 mg/ml) staining and photographed under UV light.
Nested PCR products were purified and cloned using the peqGOLD Cycle-Pure
Kit (PEQLAB) and the pCR4-Topo vector system (Invitrogen), respectively. Individual
transformed colonies were picked and directly used in a PCR reaction with the M13
forward and M13 reverse primers. Clone check PCR was carried out in 25 cycles (94°C
for 1 min, 55°C for 1 min, and 72°C for 1 min). Cloned products were then purified,
as described above, and used in cycle sequencing with the M13 PCR primers as
sequencing primers using an ABI PRISM Big Dye Terminator Cycle Sequencing Ready
Reaction Kit v.3.1 (Applied Biosystems, Warrington, United Kingdom) according
to the manufacturer's protocol and data sampling was performed on an automated
sequencer (ABI PRISM 3100 Genetic Analyser, Applied Biosystems). Sequence editing
and merging using both strand SSU and ITS sequences was done using the BioEdit
sequence Alignment Editor (version 7.0.5, http://www.mbio.ncsu.edu/BioEdit/bioedit.
html). The sequences have been deposited at GenBank (NCBI, www.ncbi.nlm.nih.gov)
under accession numbers FJ009592 - FJ009622.
Phylogenetic analysis
The SSU and ITS nuclear rDNA sequences obtained from G. intraradices, G. irregulare,
and G. geosporum were aligned using the program MAFFT (version 6.28, http://align.
bmr.kyushu-u.ac.jp/mafft/software/) resulting in a 2050 bp alignment. Phylogenetic
analyses were performed with PAUP* version 4.0b10 (Swofford 2002) as follows:
(1) DIsTANCE ANALYsIS: The BIONJ modification (Gascuel 1997) of neighbor joining
(Saitou & Nei 1987) was performed, coupled with 1000 replicates of non-parametric
bootstrapping (Felsenstein 1985), using Kimura two-parameter distances as well as
genetic distances derived from the general time-reversible model of DNA substitution,
additionally accounting for rate heterogeneity among the nucleotide sites and also
assuming a certain percentage of invariable sites (GTR+I+G; see Swofford et al. 1996 for
details). The individual model parameters were estimated using Modeltest 3.7 (Posada
& Crandall 1998).
(2) MAXIMUM LIKELIHOOD ANALYSIS: Maximum likelihood searches were performed
using PHYML 2.4 (Guidon & Gascuel 2003) under the GTR evolutionary model starting
from a tree obtained using the modified neighbor-joining algorithm BIONJ (Gascuel
1997) together with 1000 replicates of non-parametric bootstrapping.
252 ... Blaszkowski & al.
(3) MAXIMUM PARSIMONY ANALYSIS: Parsimony bootstrap values were computed
from 1000 bootstrap replicates using heuristic searches with tree bisection-reconnection
(TBR) branch swapping from starting trees obtained by subsequent addition of
sequences in random order (10 heuristic searches per bootstrap replicate, with multrees
and steepest descent options not in effect and gaps treated as missing characters).
(4) BayeEsIAN MCMC aAnatysis: Bayesian MCMC analysis was performed using
MrBayes vy. 3.1.1 (Ronquist & Huelsenbeck 2003). We ran two independent MCMC
analyses each involving two incrementally heated chains over 2 million generations
using the GIR+I+G model of nucleotide substitution and starting from random trees.
Trees were sampled every 100 generations, resulting in an overall sampling of 20 000
trees per run from which first 2000 trees were discarded (burn in). The remaining trees
sampled from each run were pooled and used to compute a majority rule consensus
tree to get estimates for the posterior probabilities. MCMC mean values of the model
parameters (obtained with Tracer v. 1.2, Rambaut & Drummond 2003) were used to
estimate branch lengths of a majority rule consensus of the sampled trees in PAUP*
(Swoftord 2002) via maximum likelihood.
Results
Both Glomus intraradices (Fics 24, 25, 29) and the fungus described below as
G. irregulare (Fics 1-4, 7, 14, 16) formed intraradical and extraradical spores
in aggregates and singly in the soil. However, marked differences mainly in (1)
the mode of spore formation, (2) spore organization in aggregates, and (3) the
phenotypic and biochemical properties of their spore wall components readily
separate the two fungi.
Phylogenetic analysis of SSU or ITS rDNA fragments, including sequences
from GenBank (data not shown), indicates that G. irregulare clusters with
G. intraradices. Although analysis using only SSU and ITS sequences obtained
in this study (Fic. 23) is better resolved, it still does not support convincingly
the obvious morphological and biochemical separation of the two fungi (Fics
1-22, 24-31). Therefore, the two fungi are described, illustrated, and compared
based on the spore morphologies and biochemical properties. Additionally, the
current distribution of G. irregulare is presented and its spores are compared
with those representing other described Glomus spp. of a similar appearance
and phylogenetic position.
Glomus irregulare Btaszk., Wubet, Renker & Buscot, sp. nov. Fics 1-23
MycoBAank No. MB 512239
Sporocarpia ignota. Sporae singulatim, gregarius in solo vel in radici efformatae. Gregaria
plerumque oblongae; 60-130 x 80-240 um; e sporis 2-4. Sporae hyalinae vel pallide luteae;
ovoideae, oblongae, vel irregulares; 60-130 x 80-240 um, raro globosae vel subglobosae;
(70-)83(-100) um diam; e alte plicatis vel profunde cavitis, 7.5-25.0 altis. Tunica sporae
e stratis tribus (strati 1-3); strato °1” caduco, glabro, hyalino, (0.5-)0.8(-1.5) um crasso;
strato “2” caduco, glabro, hyalino, (0.6-)1.8(-5.0) um crasso; strato 3” laminato, glabro,
hyalino vel pallide luteo, (1.5—)2.4(-4.4) um crasso, in solutione Melzeri rubicoso vel intenso
Glomus irregulare sp. nov. ... 253
rufuso. Hypha subtenda hyalina vel pallide lutea; recta vel recurva; fere infundibuliforma,
raro cylindrica vel colliga; (6.4-)10.8(-18.9)um lata ad basim sporae; pariete hyalino
vel pallide luteo; (1.5-)3.8(-8.5) um crasso at basim sporae, stratis 1-3 sporae continuo.
Porus apertus vel e septo continuo strati “3” sporae efformata. Vesicular-arbusculares
mycorrhizae formans.
Ho.orypvs. Poland. Szczecin, infra Plantago lanceolata L., 17 May 2006, Blaszkowski,J.
2340 (DPP).
EryMo oey. Latin, irregulare referring to the irregular shape of the fungus spores.
SPOROCARPS unknown. Spores occur mainly in aggregates inside roots or in the
soil (Fics 1-3), rarely singly in the soil (Fics 7, 14); develop blastically mainly
either at the tip of (Fic. 4) orintercalary (Fics 5-7) along dichotomously branched
coenocytic to sparsely septate hyphae continuous with mycorrhizal extraradical
hyphae (when in aggregates), rarely terminally from unbranched mycorrhizal
extraradical hyphae (when single). Hyphae bearing spores hyaline, 4.9-13.2
um wide, with a wall 1.5-3.4 um thick, composed of three layers continuous
with spore wall layers 1-3; hyphal wall layer 3 stains pale orange (6A3) to deep
red (10C8) in Melzer’s reagent (Fic. 4). The branches of the dichotomously
branched hyphae sometimes form secondary dichotomous branches bearing
next both terminal and intercalary spores. AGGREGATES usually oblong; 60-130
x 80-240 um; with 2 to 6 spores (Fics 2, 3). Spores hyaline to pale yellow
(3A3); mostly ovoid to oblong or irregular; 60-130 x 80-240 um; very rarely
globose to subglobose; (70—)83(-—100) um diam; frequently with 7.5-25.0 um
deep depressions, (Fics 1-8); spores with a single subtending hypha (Fie. 4),
occasionally with two (Fic. 7). SPORE WALL STRUCTURE composed of one wall
(Fics 4, 9-14, 16) with three layers (layers 1-3). Outermost layer 1, forming
the spore surface, semi-permanent, smooth, hyaline, (0.5-)0.8(-1.5) um thick
before disintegration (Fics 9-13, 16), frequently highly thickened at the top to
form a cap-like swelling up to 25 um high (Fic. 13). Layer 2 semi-permanent,
smooth, hyaline, (0.6—)1.8(—5.0) um thick before disintegration (F1Gs 9-13, 16).
Layers 1 and 2 usually closely adherent to each other, but separable from layer 3
and present as intact (Fics 9-13) or more or less deteriorated structures (Fics
14, 16) in most mature spores. Layer 3 laminate, smooth, hyaline to pale yellow
(3A3), (1.5-)2.4(-4.4) um thick, staining pale orange (6A3) to deep red (10C8)
in Melzer’s reagent (Fics 9-14, 16). SUBTENDING HyYPHA hyaline to pale yellow
(3A3); straight or recurvate, mostly funnel-shaped, rarely cylindrical, flared
or constricted; (6.4—)10.8(-18.9) um wide at the spore base (Fics 10, 14-16).
WALL OF SUBTENDING HYPHA hyaline to pale yellow (3A3); (1.5-)3.8(-8.5)
ttm thick at the spore base; continuous with spore wall layers 1-3 (Fries 15, 16)
in both young and most mature spores. PORE open in most mature spores (FI.
16), occasionally occluded by a curved septum (Fic. 15), (3.9-)5.9(-8.6) um
wide, continuous with the innermost lamina of spore wall layer 3. GERMINATION
STRUCTURES not observed.
254 ... Blaszkowski & al.
MYCORRHIZAL ASSOCIATIONS. The presence of mycorrhizae in field-collected
root fragments of the plant species sampled was not determined.
In single-species cultures with P. lanceolata as the host plant, G. irregulare
mycorrhizae consisted of arbuscules, vesicles, as well as intra- and extraradical
hyphae (Fics 17-22). Arbuscules were numerous and evenly dispersed along
the root fragment examined. They consisted of short trunks grown from parent
hyphae and had many branches with fine tips (Frcs 17). Vesicles usually were
ovoid to irregular, 30-120 x 60-210 um, rarely globose, (40—)91(-140) um diam
(Fics 18, 22). Their wall was thin and consisted of one layer. They frequently
occurred in aggregates widely dispersed along roots. Intraradical hyphae grew
along the root axis, were (2.2—)5.2(-9.8) um wide, straight or slightly curved, and
occasionally formed H- and/or Y-shaped branches and coils (Fics 17, 19-22).
The coils were ellipsoid, 17.5-27.5 x 37.5-82.5 u, when seen in a plane view
(Fic. 21). Extraradical hyphae were (3.2-)4.1(-—5.6) um wide and occurred in
low abundances (Fic. 22). In 0.1% trypan blue, arbuscules stained pale violet
(16A3) to reddish violet (16A7), vesicles pale violet (16A3) to deep violet
(16E8), intraradical hyphae pale violet (16A3) to greyish violet (16C6), coils -
pale violet (16A3) to light lilac (16A5), and extraradical hyphae reddish violet
(16A6-7).
PHYLOGENETIC POSITION. Five phylogenetic analyses of the nuclear SSU and
ITS rDNA sequences of spores of G. irregulare, G. intraradices, and G. geosporum
(the last serving as outgroup) indicated that, except for two G. intraradices
sequences that grouped together basal to the G. irregulare sequences, the
remaining G. intraradices sequences formed a separate cluster (Fic. 23).
SPECIMENS EXAMINED. Holotype. Poland. Szczecin, under pot-cultured P lanceolata, 17
May 2006, Blaszkowski J., 2340 (DPP); isotypes: Blaszkowski J. 2341-2355 (DPP) and
two slides at OSC.
OTHER MATERIAL EXAMINED. AFRICA, Egypt, the Giftung Island, associated with roots
of an unknown plant, 20.Aug. 2007, Blaszkowski J., 2949 and 2950 (DPP). DENMARK.
Bornholm, in roots of Ammophila arenaria, 2 Oct. 2004, Blaszkowski J., 2940-2942
(DPP). GREECE. Veriko, in the rhizosphere of Xanthium cf. spinosum L., 12 Sept.
1995, Blaszkowski J., 2943 (DPP). ITALY. Calambrone, in roots of A. arenaria, 11 Oct.
2002, Blaszkowski J. 2944 (DPP). POLAND. Hel, in roots of A. arenaria, 18 July 2000,
Btaszkowski J., unnumb. coll. (DPP); Slowinski National Park, associated with roots
of A. arenaria, 1 April 2004, Blaszkowski J., unnumb. coll. (DPP); Swinoujscie, under
Petasites spurius (Retz.) Rchb., 6 Oct. 1993, Blaszkowski J., unnumb. coll. (DPP). SPAIN.
Majorca, El Arenal, associated with roots of A. arenaria, 22 Aug. 2001, Blaszkowski J.,
a
Figs 1-8. Spores of Glomus irregulare.(DIC) 1. Intraradical spores. 2-3. Extraradical spores in
aggregates. 4. Terminal spore with 3-layered wall (swl1-3) originated from dichotomously branched
3-layered (hwl1-3) hypha. 5-6. Intercalary spores (is) with short hyphae (h) from which top spores
originated. 7. Intercalary spore with two subtending hyphae (sh). 8. Depressions (d) of spore wall.
(Figs 1, 4, 7, spores mounted in PVLG+Melzer’s reagent; Figs 2, 3, 5, 6, 8, spores mounted in
PVLG.) Bars: Figs 4-8 =10 um; Figs 2—3=20 um; Fig. 1=50 um.
Glomus irregulare sp. nov. ... 255
2.56 ... Blaszkowski & al.
2945 (DPP); Cape Salinas, associated with roots of A. arenaria, 24 Aug. 2001, Blaszkowski
J., 2946-2948 (DPP).
DISTRIBUTION AND HABITAT. Glomus irregulare spores were first found in a
field-collected rhizosphere soil of P spurius growing in maritime sand dunes
of the Baltic Sea adjacent to Swinoujscie (53°55’N, 14°14’E) in north-western
Poland in October 1993. The only other field collection of this fungus was
made in September 1995 under Xanthium spinosum colonizing dunes on the
Mediterranean Sea located near Veriko (22°35’E, 40°08’N) in Greece. Glomus
irregulare spores were also isolated from pot trap cultures with mixtures
of rhizosphere soils and Ammophila arenaria roots growing in dunes of (1)
Bornholm (55°08’N, 14°55’E), Denmark, in October 2004 (133 cultures), (2)
the Baltic Sea extending along the Stowinski National Park (54°45’N, 17°26’E)
in northern Poland in April 2004 (one culture), (3) the Baltic Sea adjacent to
Hel (54°36’N, 18°49’E) in northern Poland in July 2000 (one culture), (4) the
Mediterranean Sea near Calambrone (43°35’N, 10°18’E), Italy, in October 2002
(one culture), and (5) the Mediterranean Sea adjacent to El Arenal (39°31’N,
2°45°E) and Cape Salinas (36°19°N, 3°2°E), Majorca, Spain, in August 2001 ©
[three and one culture(s), respectively]. Additionally, G. irregulare occurred in
a trap culture with a rhizosphere soil and root fragments of an unidentified
plant growing in maritime dunes of the Giftung Island (27°11’N, 33°58’E),
Egypt, Africa, in August 2007.
In Poland, the species of arbuscular fungi co-occurring with G. irregulare in
the field were Entrophospora baltica Btaszk. et al. and G. corymbiforme Blaszk.,
and in the trap cultures Scutellospora dipurpurescens J.B. Morton & Koske. In
the field soil sample collected in Greece, G. irregulare accompanied G. xanthium
Blaszk. et al., an unrecognized Glomus sp., and S. persica (Koske & C. Walker)
C. Walker & EE. Sanders. The cultures with soils of Bornholm still contained
spores of Acaulospora mellea Spain & N.C. Schenck, Archaeospora trappei R.N.
Ames & Linderman) J.B. Morton & D. Redecker emend. Spain, G. aggregatum
N.C. Schenck & G.S. Sm. emend. Koske, G. constrictum Trappe, G. drummondii
Blaszk. & Renker, G. eburneum L.J. Kenn. et al., G. fasciculatum (Thaxt.) Gerd.
& Trappe emend. C. Walker & Koske, G. intraradices, G. mosseae, G. pustulatum
Koske et al., undescribed Glomus spp. isolates 178 and 204, Paraglomus laccatum
(Blaszk.) Renker et al., and S. dipurpurescens. In cultures with soils of Italy,
Figs 9-16. Spores of Glomus irregulare. (DIC) 9-12. Intact three layers of spore wall (swl1-3). 13.
Intact three layers of spore wall with top cap-like swell (sl) of layer 1. 14. Highly deteriorated layers
1 and 2 (swll and 2) and layer 3 of irregular spore with one subtending hypha (sh). 15. Three layers
of subtending hypha (shwl1-3) occluded by curved septum (s). 16. Subtending hyphal wall layers
1-3 (shwl1-3) continuous with spore wall layers 1-3 (swl1-3).
(Figs 11-14, 16, spores mounted in PVLG+Melzer’s reagent; Figs 10, 15, spores mounted in
PVLG.) Bar =10 um.
Glomus irregulare sp. nov. ... 257
258 ... Blaszkowski & al.
Se
Figs 17-22. Mycorrhizae of Glomus irregulare in roots of Plantago lanceolata stained in 0.1%
trypan blue. (bright field). 17. Arbuscule (arb) with trunk (tr). 18. Vesicles (ves). 19. H-shaped
branch (Hb). 20. Y-shaped branch (Yb). 21. Coils (c). 22. Vesicles (ves), straight intraradical hypha
(ih), extraradical hyphae (eh), and young spore (sp).
Bars: Figs 17, 19-21 =10 um; Figs 1, 18, 22=50 pm.
spores of G. aurantium Blaszk. et al. and an undescribed Glomus sp. were also
found. Apart from G. irregulare, the arbuscular fungi sporulating in Majorca’s
cultures also were G. aurantium, G. constrictum, Pacispora scintillans (S.L. Rose
& Trappe) Sieverd. & Oehl, and S. calospora (T.H. Nicolson& Gerd.) C. Walker
Glomus irregulare sp. nov. ... 259
& FE. Sanders. In the culture with the soil of the Giftung Island, spores of G.
drummondii were still found.
The soil chemical properties of the dunes adjacent to Swinoujécie ranged:
Pei3.o-6.7; NO, (mg L*), 20-72; P, 5-12; K, 2-26; Mg, 10-41; Na, 4-23; Cl,
15-25; KCl, 0.1-0.6; organic C (%), 0.1-1.1 (Blaszkowski 1995).
Discussion
The unique properties of Glomus irregulare are (1) the frequent formation
of irregular spores with deep wall depressions and apical cap-like swellings
of their outermost wall layer 1 (Fics 2, 5, 6, 8, 13, 14), (2) the production of
spores mainly in oblong aggregates containing spores arising either from the
tip of or intercalary along dichotomously branched hyphae continuous with
mycorrhizal extraradical hyphae (Fic. 2), and (3) the intense reactivity of the
laminate innermost spore wall layer in Melzer’s reagent (Fics 4, 7, 11-14, 16).
This fungus is additionally distinguished by the tendency to form spores mainly
inside roots of its host plant (Fic. 1).
Our molecular analyses (Fic. 23, see above) show G. intraradices as the
fungus most closely related to G. irregulare, although the analyses did not
prove conclusively their separateness. In this study, the G. irregulare and
G. intraradices spores used in phylogenetic analyses and to compare their
morphologies and biochemical properties came from single-species pot
cultures derived from pot trap cultures containing rhizosphere soils and
A. arenaria roots from the Bornholm maritime dunes. Thus, the morphological,
_ biochemical, and molecular divergences did not result from the influence
_ of heterogeneous environments. Morton & Benny (1990b) found that spore
_ characteristics of G. intraradices clonal populations remained consistent, even
though they originated from highly differing environments. Lloyd-Macgilp et
al. (1996), who compared the genetic diversity of arbuscular mycorrhizal fungi
| between and within different continents, noted that diversity did not increase
| with increasing geographical distance. Hence, the differences in the mode of
| formation of spores, their organization in aggregates, and the phenotypic and
| biochemical properties of components of their spore wall markedly separating
| the two fungi (see below comparisons and Fics 1-22, 24-31) are rather innate
_ and, thereby, sufficiently important to support G. irregulare as a new taxon.
Seven properties separate G. irregulare from G. intraradices. First, whereas
_ the distinctive property of G. irregulare is the formation of aggregates with
| spores arising both terminally and intercalary (Fics 2-8), the G. intraradices
aggregates contain only terminal spores (Fic. 24; Blaszkowski, pers. observ.).
Moreover, the intercalary mode of spore formation in G. irregulare usually
| causes the aggregates to be oblong and compact (Fics 2, 5, 6) rather than globose
to ovoid (or irregular and loose) as in G. intraradices (Fic. 24; Blaszkowski
260 ... Blaszkowski & al.
2003, Morton 2002). Second, the darkest G. irregulare spores [pale yellow
(3A3); Fias 2, 5, 6, 8, 9, 10, and 15] are much lighter-coloured than the darkest
G. intraradices spores [yellow brown to grey brown: Morton 2002, Schenck &
Smith 1982; greyish yellow (2B5): Blaszkowski 2003], which still frequently
have a greenish tint (Fic. 24; Blaszkowski 2003, Morton 2002, Schenck & Smith
1982; vs. no such a tint in G. irregulare spores). Third, most G. irregulare spores
are irregular in shape and have distinctive wall depressions and apical cap-like
swellings (Fics 1-3, 5, 6, 8, 13, 14). In contrast, almost all G. intraradices spores
are globose to subglobose, and their wall has neither depressions nor swelling
at the spore apex (Fics 24-31). Fourth, the two outer wall layers of G. irregulare
spores are semi-permanent and usually intact or only slightly deteriorated in
mature specimens (Fics 4, 9, 10, 11-14, 16), whereas those of G. intraradices
are short-lived and frequently highly decomposed or completely sloughed at
maturity (Fics 30, 31). The mucilaginous nature of the outermost wall layer of
G. intraradices spores roughens their surface (Fics 26, 27, 29, 30). In contrast,
the G. irregulare spore surface usually is smooth (Fics 4, 6, 8-16). Fifth, the
laminate layer of G. irregulare spores consists of generally inseparable, thin (ca.
0.5 um thick) laminae (Fics 4-14, 16), while the G. intraradices laminate spore
layer comprises frequently separating, 0.5-1 um thick, sublayers (Fics 26, 28,
31; Blaszkowski 2003, Morton 2002, Schenck & Smith 1982). The laminate
layer also is much thinner (1.5-4.4 um) in G. irregulare than in G. intraradices
(3.2-12.0 um). Sixth, the staining spore wall layer of G. irregulare in Melzer’s
reagent is only the laminate innermost layer (Fics 4, 7, 11-14, 16), whereas
the reactive wall layer of G. intraradices spores in Melzer’s is the mucilaginous
outermost one (Fics 26-30; Blaszkowski 2003, Stiirmer & Morton 1997).
Finally, G. irregulare distinctively tends to hide its spores inside roots (Fic.
1) and only rarely produces extraradical spores, especially in field conditions.
For example, ca. 160 field-collected mixtures of rhizosphere soils and roots of |
plants colonizing the Bornholm maritime dunes did not contain G. irregulare )
spores, but they occurred in over 80% of pot trap cultures of these mixtures
after greenhouse cultivation (Blaszkowski, pers. observ.). Moreover, even
in both trap and single-species cultures, G. irregulare did not begin to form
extraradical spores until the third or the fourth cultivation cycle, although
its numerous intraradical spores were still frequently visible in early control
:
:
|
Fig. 23. BIONJ tree based on an alignment of nuclear SSU and ITS rDNA sequences obtained
from Glomus irregulare and Glomus intraradices spores. A total of 2050 alignment positions were
used for tree construction. The tree was rooted with Glomus geosporum. Branch support values
(only values exceeding 50% are given) on branches refer to BIONJ bootstrap, BIONJ bootstrap
(using GTR+I+G distances), Phyml bootstrap, maximum parsimony bootstrap, and posterior |
probabilities as estimated by Bayesian MCMC (GTR+I+G model of DNA substitution), respectively. |
All sequences were obtained in this study. ;
Glomus irregulare sp. nov. ... 261
G. irregulare 04
' G. irregulare 06
G, irregulare 08
G. irregulare 01
G. irregulare 07
G. irregulare 02
G. irregulare 05
G. irregulare 03
~ G. irregulare 14
100/ 60/ 94/ 99/ -
~~ G, itregulare 10
G. irreguiare 12
G. irregulare 11
86/ -/-/97/- :
G. irregulare 13
iG irregulare 09
G. intraradices 02
59/ 58/ 81/ 99 74
G. intraradices 05
G. intraradices 03
G. intraradices 08
100/ 100/ 100/ 100/ 100 G. intraradices 04
G. intraradices 12
G. intraradices 14
G, intraradices 15
G. intraradices 09
G. intraradices 11
G. intraradices 13
100/ 97/ 98/ 99/ 100
G. intraradices 10
G. intraradices 07
G. geosporum 01
G. geosporum 03
G. geosporum 04
Ria? ; G. geosporum 02
———~ (0.005 substitutions/site cea
262 ... Blaszkowski & al.
harvests of the first cycle cultures. In contrast, G. intraradices usually sporulated
in the field and always formed numerous both intra- and extraradical spores
during the first cultivation cycle in single- and multi-species cultures (Fic. 24;
Blaszkowski, pers. observ.).
Another species of arbuscular mycorrhizal fungi of the genus Glomus
morphologically resembling G. irregulare is G. aggregatum. This fungus also
produces spores both in aggregates within roots and the soil and singly in the
soil (Blaszkowski 1991, 2003; Koske 1985, Schenck & Smith 1982). Moreover,
its spores are similar in size, somewhat in shape, and in having a 3-layered
wall with two sloughing, hyaline outer layers and a laminate, yellow-coloured
innermost layer. Although G. aggregatum frequently forms irregular spores, they
have neither deep depressions nor apical cap-like swellings that characterize
G. irregulare spores. Spores of G. irregulare with depressions also occur inside
roots, suggesting that depressions are a genetically conserved character of this
fungus rather than influenced by external conditions during spore development,
e.g., caused by lack of space for neighbouring spores.
Apart from forming spores in aggregates and singly in the soil, G. aggregatum
also produces new spores by internal proliferation within their parent spores
(Btaszkowski 2003, Koske 1985), a phenomenon not found in G. irregulare.
Additionally, G. irregulare spores are much lighter-coloured [hyaline to pale
yellow (3A3)] than those of G. aggregatum [pale yellow to orange brown: Koske
1985; pastel yellow (2A4) to yellowish brown (5D8): Blaszkowski 2003].
The results of our phylogenetic analysis presented in Fic. 23 and analysis of
two datasets based on SSU fragment and 5.8S together with the highly variable
ITS2 region including related sequences available from GenBank also indicated
that G. irregulare is closely related to the G. intraradices species complex (data
not shown). Our data also showed that two G. intraradices sequences formed
a separate subgroup associated with those of G. irregulare. This incomplete
agreement between the evident differences in morphological properties of
spores of G. irregulare and G. intraradices and the results of our phylogenetic
analyses might be due to the highly conserved molecular markers used in this
study. Hence, we suggest the use of additional or other genetic markers in
further studies to explain the inconsistency revealed here.
Figs 24-31. Spores of Glomus intraradices. 24. Loose aggregate of intact spores. (bright field)
25. Intraradical spores. 26-28. Spore wall layers 1-3 (swl1-3). 29. Spore wall layers 1-3 (swl1-3)
continuous with subtending hyphal wall layers 1-3 (shwl1-3). 30. Spore wall layers 1-3 (swl1-3);
note the highly deteriorated swl1. 31. Spore wall layers 2 and 3 (swl2 and 3); layer 1 is completely —
sloughed in this spore.
(Fig. 24 bright field, spores in lactic acid. Figs. 25-31 DIC, spores mounted in PVLG+Melzer’s
reagent Fig. 31, spores mounted in PVLG)
Bars: Figs 26-31=10 um; Fig. 25=50 um; Fig. 24=100 um.
263
Glomus irregulare sp. nov. .
264 ... Blaszkowski & al.
As concluded above, G. irregulare is most closely related in molecular
phylogenetic analyses to G. intraradices, a member of the subclade Glomus group
Ab, also comprising G. clarum T.H. Nicolson& N.C. Schenck, G. coremioides
(Berk. & Broome) D. Redecker & J.B. Morton, G. fasciculatum, G. manihotis
R.H. Howeler et al., G. proliferum Dalpé & Declerck, G. sinuosum (Gerd. & B.K.
Bakshi) R.T. Almeida & N.C. Schenck, and G. vesiculiferum (Thaxt.) Gerd. &
Trappe (Schwarzott et al. 2001). Morphologically, however, G. irregulare more
or less resembles only G. fasciculatum, G. proliferum, and G. vesiculiferum,
fungi also producing spores in conglomerations. i
In contrast to the primarily intercalary formation of irregular spores in G.
irregulare, G. fasciculatum spores always arise terminally and are globose to
subglobose (Blaszkowski 2003, Walker & Koske 1987). Moreover, the only spore
wall component shared by the two species is their laminate layer. However,
the laminate layer is the innermost structure of the G. irregulare spore wall,
whereas in G. fasciculatum the laminae occur in the middle of the 3-layered
spore wall. In the latter species, the two other spore wall layers are a permanent
outermost layer and a flexible innermost one, whereas the laminate layer does’
not differentiate at all in G. irregulare.
Although G. irregulare and G. proliferum form intercalary and terminal spores
in the soil and roots, G. proliferum spores are usually globose to subglobose
(vs. mainly ovoid to irregular in the former fungus), markedly smaller [(22-
)40-66(-76) um diam vs. (70-)83(-100) um diam when globose], and have
different spore wall phenotypic and biochemical properties (Blaszkowski
2003, Declerck et al. 2000). Phenotypically, these fungi link only their laminate
innermost layer. However, this layer stains in Melzer’s reagent in G. irregulare,
and is nonreactive in G. proliferum.
The globose spores of G. irregulare and G. vesiculiferum are similar in
colour and size, but G. vesiculiferum is unique in that its spores are grouped
in sporocarps with a peridium-like layer consisting of thin-walled, ellipsoid to
broadly clavate vesicles (Gerdemann & Trappe 1974). In contrast, G. irregulare
spores occur in loose aggregates with no peridium and/or singly in the soil.
Additionally, the G. vesiculiferum spore wall is 2-layered, not 3-layered as in G.
irregulare. Unfortunately, the reactivity of G. vesiculiferum spores in Melzer’s
reagent is unknown.
Although G. irregulare was identified in only 143 of ca. 5000 mixtures of |
rhizosphere soils and roots examined by the first author of this paper during
the last 25 years (of which 133 mixtures came from Bornholm), the presence
of this fungus in different regions of Europe, Africa and in Australia (Dr. C.
Walker, Royal Botanic Garden Edinburgh; pers. comm.) suggests it is widely
distributed in the world. The rare finding of G. irregulare probably results
from the tendency to hide its spores inside roots and the exceptionally rare
Glomus irregulare sp. nov. ... 265
production of extraradical spores. For example, in the field soil samples
collected in Swinoujscie (Poland) and Veriko (Greece), in which extraradical
spores were found, their abundance in 100 g dry soil was very low, i. e., 11 and
2, respectively.
Acknowledgements
This study was supported in part by The Committee of Scientific Researches, grants
no. 2 PO4C 041 28 and 164/N-COST/2008/0. Many thanks to Sabine Jarzombski and
Bettina Schlitt for their help in cloning and sequencing. Thanks are due to Dr. Gabor M.
Kovacs, Department of Plant Anatomy, Eotvos Lorand University, Budapest, Hungary
and Priv. Doz. Dr. habil Ewald Sieverding, Institute of Plant Production and Agroecology
in the Tropics and Subtropics, University of Hohenheim, Germany for serving as pre-
submission reviewers.
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closed pot cultures of arbuscular mycorrhizal fungi. Agricultural Science in Finland 3, 233-
240.
Walker C, Vestberg M, Demircik F, Stockinger H, Saito M, Sawaki H, Nishmura J, Schtifler A. 2007.
Molecular phylogeny and new taxa in the Archaeosporales (Glomeromycota): Ambispora fennica
gen. sp. nov., Ambisporaceae fam. nov., and emendation of Archaeospora and Archaeosporaceae.
Mycological Research 111: 137-13.
Walker C, Vestberg M, Schiifler A. 2007. Nomenclatural clarifications in Glomeromycota.
Mycological Research 111: 253-255.
Wubet T, Weifs M, Kottke I, Teketay D, Oberwinkler E 2006. Phylogenetic analysis of nearly
complete SSU rDNA sequences reveals that the endangered African Pencil Cedar, Juniperus
procera Hochst. ex Endl., is associated with distinct Glomus taxa. Mycological Research 110:
1059-1069.
MYCOTAXON
Volume 106, pp. 269-272 October-December 2008
Calvatia sculpta, a striking puffball occurring on
Brazilian sand dunes
IURI GOULART BASEIA ! & FRANCISCO DIEGO CALONGE 2
baseia@pesquisador.cnpq.br
'Depto. Botanica, Ecologia e Zoologia
Universidade Federal do Rio Grande do Norte
Campus Universitario, CEP: 59072-970, Natal, RN, Brazil
calonge@rjb.csis.es
? Real Jardin Botanico, CSIC
Plaza de Murillo 2, 28014. Madrid, Spain
Abstract — This North American species was discovered for the first time in South
America, growing on sand dunes associated with native tree roots. A description,
comments, illustration of the basidiome and SEM-photo of the spores are given.
Key words — Lycoperdaceae, gasteromycetes, neotropics, taxonomy, chorology
Introduction
H. W. Harkness described Lycoperdon sculptum in 1885 from specimens
collected in the Sierra Nevada Mountains in California, and in 1904 Lloyd
transferred the species to the genus Calvatia. This North American species has
avery restricted geographic distribution and is considered uncommon by some
mycologists (Arora 1986). It is quite unique, owing to its basidiomata with
surfaces covered by various pyramid-like structures. It can easily be confused
with Calbovista subsculpta Morse ex M.T. Seidl, which exhibits felty warts and a
root-like soil attachment. In some cases it can also be confused with immature
specimens of Amanita magniverrucata Thiers & Ammirati. A classification in
_ the Calvatia complex was proposed by Lange (1993).
Material and methods
Field expeditions were made in June 2007 in an Atlantic forest remnant in
| Parque Estadual das Dunas in Natal (05°46’ S, 35° 12 W), Rio Grande do Norte
| state. The specimens examined are deposited in the Mycological Herbarium
of the Botany, Ecology and Zoology Department in Natal, Brazil (UFRN).
270 ... Baseia & Calonge
Macroscopic characters were described based on observations of fresh and dried
material. Free-hand sections of gleba and peridium were mounted in water, 3%
(w/v) KOH, and 2% cotton blue lactophenol on glass slides and observed under
a light microscope (LM). Forty randomly selected basidiospores were measured
under the LM at 1000x magnification. Spores from the gleba were dusted onto
specimen holders attached with double-sided carbon adhesive tape and then
coated with up to 15 angstroms of gold-palladium on an Ion Sputter Coater
to prepare for scanning electron microscopy (SEM). Colors were determined
according to Kornerup & Wanscher (1978).
Results
Calvatia sculpta (Harkn.) Lloyd, Myc. Writ. 1: 203. 1904. Figs. 1-2
Basionym: Lycoperdon sculptum Harkn., Calif. Acad. Sci. Bull. 1(3): 160. 1885.
Basidiomata epigeous, subglobose, 7-9 cm broad, white. Exoperidium thick,
rimose and cracking into pyramidal warts horizontally grooved by parallel lines
(Fig 1). Endoperidium thin, fragile. Sterile base present occupying one third of
the basidioma, chambered, yellowish white (4A2), lacking a rooted base. Gleba
pulverulent, pale yellowish (4A3). Capillitial threads 4-9 um broad, branched,
septate, pitted, light brown. Basidiospores globose, 3.5-4 um broad, minutely
verrucose under SEM, pedicel short (Fig. 2), light brown.
KNOWN DISTRIBUTION: United States of America (Harkness 1885, Coker & Couch 1928,
Zeller 1964, Arora 1986)
MATERIAL EXAMINED: BRAZIL, Rio Grande do Norte, Natal, Parque Estadual das
Dunas, Natal, on sandy soil, Baseia, 13-VI-2007 (UFRN-fungos 612).
Notes: Basidiospore sizes vary greatly in C. sculpta, with spores ranging in
size from 3.5-4.0 um to 7.2-9.5 um diam in specimens collected in the USA
(Kreisel, pers. comm.). The material collected in Brazil has spores in the lowest
margin (3.5-4.0 um).
Discussion
Calvatia sculpta is a North American species that is normally associated with
conifers in the inland mountains of California, Oregon, Washington, and |
Idaho (Zeller & Smith 1964). It appears restricted in range and has yet to be
reported from the Pacific coast (Arora 1986). The discovery of this species
|
in a dune biome in the Northeast of Brazil associated with native tree roots —
(Eugenia brasiliensis Lam., Myrtaceae) challenges hypotheses by Mueller et |
al. (2007) regarding its ecology and geographic distribution. One hypothesis _
proposes that C. sculpta is an ancient species that already existed before the —
Americas drifted apart. A second suggests that a dispersal agent (likely an
Calvatia sculpta in Brazil ... 271
Fic. 1. Calvatia sculpta, general view of the basidioma.
Fic. 2. Calvatia sculpta. details of the basidiospores under SEM.
anthropological action) introduced this organism into new biomes where it
adapted by associating itself with other plant species. A third hypothesis is
that there are at least two distinct species that share similar morphological
features and correspond to cryptic species. Molecular studies are needed to test
the hypotheses and help determine the true relationships between the North
American and South American C. sculpta populations.
272 ... Baseia & Calonge
The systematics of Calvatia have been modified during the past decade
based on morphological characters. In 1992, Kreisel proposed sect. Sculpta
to accommodate C. sculpta; two years later the same author (Kreisel 1994)
combined sections Cretacea and Sculpta based mainly on characters published
by Lange (1990, 1993, 1994), thus placing C. sculpta close to C. arctica Ferd. &
Winge.
Acknowledgments
This research is supported by the Conselho Nacional de Desenvolvimento Cientifico e
Tecnoldgico (CNPq), Brazilian agency. The authors would like to thank Prof. H. Kreisel,
Prof. G. Moreno and Dr. Maria Alice Neves for useful comments and Tereza Cristina de
Oliveira Galvao for the drawing.
References
Arora D. 1986. Mushrooms demystified: A comprehensive guide.to the fleshy fungi. Ten Speed
Préss,,CA,212 pp.
Coker WC, Couch JN. 1928. The gasteromycetes of the Eastern United States and Canada, Chapel
Hill, 201 pp.
Harkness HW. 1885. Fungi of the Pacific Coast. Bull. Calif. Acad. Sci. 1(3): 159-177.
Kornerup A, Wanscher JH. 1978. Methuen handbook of colours. Third edition. Eyre Methuen.
London. 252 pp.
Kreisel H. 1992. An emendation and preliminary survey of the genus Calvatia (Gasteromycetidae).
Persoonia 14: 431-439.
Kreisel H. 1994. Studies in the Calvatia complex (Basidiomycetes) 2. Feddes Repertorium 105:
369-376.
Lange M 1990. Arctic Gasteromycetes II. Calvatia in Greenland, Svalbard and Iceland. Nordic J.
Bot., 9: 525-546.
Lange M. 1993. Classification in the Calvatia group Blyttia 3-4: 141-144.
Lange M.1994. Calvatia subcretacea, a synonym of C. arctica. Sydowia 46: 126-129.
Lloyd CH. 1904. Mycological notes, no. 18. Mycol. Writ. 1: 189-204.
Mueller GM, Schmit JP, Leacock PR, Buyck B, Cifuentes J, Desjardin DE, Halling RE, Hjortstam K,
Iturriaga T, Larsson KH, Lodge DJ, May TW, Minter D, Rajchenberg M, Redhead SA, Ryvarden
L, Trappe JM, Watling R, Wu Q. 2007. Global diversity and distribution of macrofungi.
Biodiversity and Conservation 16(1): 37-48.
Zeller SM, Smith AH. 1964. The genus Calvatia in North America. Lloydia 27: 148-180.
MY COTA XON
Volume 106, pp. 273-288 October-December 2008
Selected species of Mucorales from soil contaminated
with toxic metals in Sao Paulo State, Brazil
JOSE IVANILDO DE SOUZA’, IRACEMA HELENA SCHOENLEIN-CRUSIUS?
& LuisA HELENA DOS SANTOS OLIVEIRA?
1- jisouza@yahoo.com.br 2- iracema@crusius.com.br
3- luisahelena_oliveira@yahoo.com.br
Instituto de Botanica, Segao de Micologia e Liquenologia
Av. Miguel Stefano 3687, 04301-902, Sao Paulo - SP, Brazil
NORBERTO CARLOS SCHOENLEIN
norbertocs@cy.com.br
Universidade Guarulhos, Centro de Pés-Graduacdo, Pesquisa e Extensao
Praca Teresa Cristina n° 1, 13067-067, Guarulhos - SP, Brazil
SANDRA FARTO BOTELHO TRUFEM
strufem@cyberspace.com.br
Universidade Sao Marcos
Rua Clovis Bueno de Azevedo 176, 04266-040, Sao Paulo - SP. Brazil
Abstract — Eleven taxa of Mucorales were isolated from soil-plates made with malt
agar amended with 0, 100, 200, 500 or 1,000 mg/L of Pb(NO,), or Zn(NO,), and soil
collected in an area contaminated with chemicals. The taxa isolated (Mucor bainieri, M.
circinelloides f. circinelloides, M. circinelloides f. lusitanicus, M. circinelloides f. janssenii,
M. hiemalis f. hiemalis, M. hiemalis f. luteus, M. racemosus f. racemosus, Zygorhynchus
moelleri, Absidia cylindrospora v. cylindrospora, Rhizopus arrhizus v. arrhizus and
Cunninghamella phaeospora v. phaeospora) are described, discussed, and illustrated.
This is a second world record for Mucor bainieri, previously isolated only once in India,
and the first record this species from Brazil.
Key words — soil fungi, soil pollution, Mucoromycotina, zygomycete
Introduction
Most Brazilian studies on Mucorales (Mucoromycotina) have been carried out
in the northeast, north and southeast regions of the Brazil. These fungi have
been isolated from herbivore dung (Batista 1948, Alves et al. 2002, Santiago &
Cavalcanti 2007, Santiago 2008, Santiago et al. 2008), as contaminants in other
274 ... de Souza & al.
fungal cultures (Batista & Vital 1955) and corn flour (Ribeiro et al. 2003), from
sugar cane and rice cultivated soils and other kinds of soils (Upadhyay 1967,
1969, 1970, 1973, Santos et al. 1989) in Pernambuco State, decaying flowers
and soil (Dantas 1953ab) and copper-mining soils (Santiago & Souza-Motta
2006) in Bahia State and soil in the States of Maranhao (Lira 1971) and Para
(Pfenning 1993).
In southeast, Sao Paulo State, another sequence of studies was performed
on Mucorales from soil (Trufem 1978, Trufem 198la-—c, Antunes et al. 1993,
Santos et al. 1998, Ruegger et al. 2002, Viriato 2003, Tauk-Tornisielo et al.
2005), herbivore dung (Trufem 1984, Trufem & Viriato 1985, Viriato & Trufem
1985a,b, Viriato 2003), water, soil and leaves (Schoenlein-Crusius & Milanez
1989, 1997, Wellbaum et al. 1999), soil polluted with bird dung (Schoenlein-
Crusius et al. 1996), air (Schoenlein-Crusius et al. 2001), and soil and leaves
from areas damaged by industrial pollution (Schoenlein-Crusius et al. 2006).
Efforts have been made to evaluate the state of knowledge of Brazilian
macro- and microfungi, including Mucorales, from Cerrado reserves (Brazilian
Savanna) in Sao Paulo State (Milanez et al. 1997), Atlantic Rain Forest reserves.
in Pernambuco State (Maia et al. 2006), areas of Sao Paulo State, of the northeast
and its semi-arid regions (Trufem 1999, Trufem et al. 2006), microfungi
(including Mucorales) from several regions and biomes (Pfenning 1997) and
Mucorales of medical importance (Lacaz et al. 1998).
Herein we report on the distribution of additional Brazilian Mucorales.
Material and methods
Soil samples were collected up to 10 cm in depth at five sites in Santa Gertrudes
Municipality, Sao Paulo State, Brazil, and each sample was a composite of three sub-
samples collected in adjacent points (de Souza 2006). Four of the sites were located in an
interdicted area at Fazenda Itaqui drainage basin near Santa Gertrudes glazed ceramics
industrial complex and km 168 of Washington Luiz highway, where the industries
discharged ceramic wastes containing toxic metals (i.e. Pb and Zn) and other chemicals
that has contaminated the soil, groundwater and surface water (Bonacin Silva 2001). The
use this area is banned by the environmental authorities until conclusion of actions and
studies for its recovery. The fifth site was in the urban area of Santa Gertrudes (control
site). Soil suspensions (10% w/v) were prepared using distilled sterile water transferred
to 250 mL Erlenmeyer flasks, which were shaken 1.5 h at 70 rpm. In order to prepare
soil-plates (Warcup 1950), 1 mL of the soil suspension was spread over the surface of a
Petri dish containing MA (malt agar 2% w/v) previously amended with 0 (control), or
100, 200, 500 or 1,000 mg/L of Pb(NO,), or Zn(NO,),.6H,O (based in Arnebrant et al.
1987). The assays were carried out in triplicate and the Petri dishes incubated 7 days at
20-23 °C. The colonies were transferred to PDA (potato dextrose agar) and incubated 7
days at 20-23 °C, then transferred to SMA (synthetic Mucor agar, Hesseltine 1954) and
incubated in the dark 5-7 days at 25 °C, and when necessary at 20, 30, 37, 40 or 45 °C.
Mucorales from contaminated soils (Brazil) ... 275
Slides were prepared with distilled water plus glycerol (2% v/v) or PVLG resin (Morton
et al. 1993) at 50% (w/v). Twenty measurements were made for each fungal structure. The
identifications were performed in accordance with the literature on Absidia (Hesseltine
& Ellis 1964, Schipper 1990), Cunninghamella (Baijal & Mehrotra 1980, Lunn & Shipton
1983, Zheng & Chen 2001), Mucor (Benjamin & Mehrotra 1963, Schipper 1973, 1976),
Rhizopus (Schipper 1984) and Zygorhynchus (Hesseltine et al. 1959, Schipper 1986). The
cultures were preserved by immersion in sterile distilled water, lyophilized, numbered,
and stored in the fungi culture collection of the Secao de Micologia e Liquenologia,
Instituto de Botanica, in Sao Paulo City, Sao Paulo State, Brazil.
Results and discussion
Mucor bainieri B.S. Mehrotra & Baijal, Aliso 5: 237. 1963. PLATE 1
SPECIMEN EXAMINED: Brazil, Sao Paulo State, Santa Gertrudes, Fazenda Itaqui drainage
basin, contaminated area near ceramic tile industries, 22°27'46”S 47°32’29” W, soil under
grass, J.I. de Souza & N.C. Schoenlein, 30.V1.2003, No. 2033.
Colonies 15-30 days old on SMA at 20 and 25 °C, producing a strong baker's
yeast odor, obverse with thin mycelium, pale beige dotted in chestnut, or
whitish with or without a yellowish cream margin, reverse yellowish cream, pale
beige to orange beige, 71 x 68 mm in size, up to 7 mm high; sporangiophores
sympodially branched, 57-1,560 um long, 3.2-11.7 um in diameter at base and
2.7-9.2 tum in diameter at apex; sporangia globose, slightly applanate, reddish
amber to chestnut, sporangial wall deliquescent, 24-61 x 25-67 um; columellae
ellipsoidal, obovoid, pyriform, some globose, hyaline green to slightly greyish,
collars present or absent, 11.6-40 x 13.2-45 um; sporangiospores ellipsoidal,
flattened at one side, hyaline green, 4.2-9.7 x 2.2-5 um; azygophores branched
or unbranched, sometimes on the same branch with sporangiophores, 5-12.5
lm in diameter; azygosporangia applanate, reddish amber to dark brown, some
moss green (young), on azygophore apices, laterally located on some aerial or
submerged hyphae, present in single-spore cultures, 26-62 x 28-75 um, spines
isolated or on stellar warts (starfish-like) up to 5 um high; azygospores from
azygosporangia not observed; chlamydospores isolated, catenate, apical or
intercalate, hyaline to green, containing olive green to orange droplets, 7.2-26
x 8-16.4 um; submerged mycelium containing olive green to orange droplets.
Notes. Isolated on MA with 200 mg of Pb(NO,),. After 5 days on SMA at 30
°C both normal growth and sporulation; after 5 days at 37 °C no growth but the
culture grew when placed again at 25 °C. This rare species of Mucor was initially
recorded from forest soil in Uttar Pradesh, India (Benjamin & Mehrotra 1963).
The strain No. 2033, which corresponds to the first record of M. bainieri from
Brazil, constitutes only the second world record. Schipper (1976) supplied
another description of the original strain (CBS 293.63) showing results of
influence of light, media and inter-specific mating.
276 ... de Souza & al.
PLATE 1. Mucor bainieri. F1G. 1. Branched azygophore with terminal azygosporangium. FIGS. 2, 3.
Azygosporangia. FIG. 4. Azygosporangium with stellar warts, in the submerged mycelium. Fic. 5.
Sporangium. FIGs. 6-8. Columellae. Fic. 9. Sporangiospores. Bars = 10 um.
Mucor circinelloides Tiegh., Annis. Sci. Nat. Bot., Sér. 6, 1: 94. 1875.
f. circinelloides PLATE 2 (Fics. 1-5)
SPECIMEN EXAMINED: Brazil, Sao Paulo State, Santa Gertrudes, urban area, 22°27712”S
47°32°39°W, soil from a public garden, J.J. de Souza & N.C. Schoenlein, 30.V1.2003, No.
2034.
Colonies 5 days old on SMA at 25 °C, obverse beige dotted with chestnut,
reverse greyish beige, pale orange at inoculum region, 90 mm in diameter, up to
6 mm high; sporangiophores sympodially branched, curved to almost circinate
the short or straight the long, 22.5-1,010 um long, 6.3-16.3 um in diameter at
base and 5-12.5 um in diameter at apex; sporangia globose, slightly applanate,
greenish chestnut to greenish golden yellow, sporangial wall deliquescent or
persistent, 28-53 x 28-58 um; columellae ellipsoidal, obovoid, or globose
slightly applanate, mostly with collars, hyaline to brownish, 10.5-43 x 11.5-44
[Lm; sporangiospores globose, ovoid or ellipsoidal, smooth with homogeneous
Mucorales from contaminated soils (Brazil) ... 277
contents, olive green to brownish, 4.3-12.3 x 3.3-8.8 um; chlamydospores
apical, intercalate or catenate, thick walled, granular, with olive green to golden
yellow contents, 18-41 x 13-32 um; submerged mycelium three times as wide
as the aerial hyphae and replete with olive green contents; zygosporangia not
observed.
Notes. Isolated on MA with 100 mg of Pb(NO,),. After 7 days on SMA at 37
°C normal growth and reduced sporulation; after 7 days at 40 °C, no growth.
The form circinelloides was isolated from soil (Schoenlein-Crusius et al. 1996,
2006) and herbivore dung (Alves et al. 2002, Santiago 2008); M. circinelloides
was isolated from soil (Trufem 1978, Antunes et al. 1993, Ruegger et al. 2002),
herbivore dung (Trufem & Viriato 1985) and leaves (Schoenlein-Crusius &
Milanez 1989).
Mucor circinelloides f. lusitanicus (Bruderl.) Schipper,
Stud. Mycol. 12: 9. 1976. PLATE 2 (FIGS. 6-8)
SPECIMEN EXAMINED: Brazil, Sao Paulo State, Santa Gertrudes, urban area, 22°27712”S
47°32’39°W, soil from a public garden, J.I. de Souza & N.C. Schoenlein, 30.V1.2003, No.
2035.
Colonies 6 days old on SMA at 25 °C, obverse powdery grey or greenish
chestnut, with or without a whitish to beige margin, pale orange at the inoculum
site, reverse pale beige, slightly grey, 78 mm in diameter, 2 mm high centrally
and 1 mm high on the remainder of the colony; sporangiophores sympodially
branched, short or long, slightly curved, 9.2-1,550 um long, 4.5-17.5 um in
diameter at base and 3.5-12.5 um in diameter at apex, sometimes forming
a flaccid turf up to 5 mm high; sporangia globose, applanate, golden yellow,
greenish chestnut to almost black, sporangial wall deliquescent or persistent,
29-78 x 30-85 um; columellae globose, slightly applanate, with large basal
membranes or smooth and without collars, hyaline to light grey, with or
without scattered granular contents, 9.2-48 x 10.2-50 um; sporangiospores
ellipsoidal, ovoid, or less frequently kidney- or drop-shaped, with a granular
appearance, olive green, slightly pale chestnut spotted, 3.2-10.2 x 2.7-5.7
tum; chlamydospores apical, thick walled, full of olive green contents, 10.2-31
x 9.2-25 um; submerged mycelium with greenish golden yellow contents;
zygosporangia not observed.
Notes. Isolated on MA with 200 mg of Pb(NO,),. After 7 days on SMA at 37 °C
reduced growth and no sporulation; after 7 days at 40 °C no growth, however,
growth commenced again when the colonies were placed at 25 °C. This form of
M. circinelloides was isolated from herbivore dung (Alves et al. 2002, Santiago
2008).
278 ... de Souza & al.
PLATE 2. FIGS. 1-5. Mucor circinelloides f. circinelloides: 1. Short sympodial branches. 2. Sporangium
on short sporangiophore. 3, 5. Columellae on short and long sporangiophores, respectively.
4. Sporangiospores. FIGs. 6-8. M. circinelloides f. lusitanicus: 6. Sporangium. 7. Columella.
8. Sporangiospores. FIGS. 9-12. M. circinelloides f. janssenii: 9. Short sympodial branches.
10. Sporangium on short sporangiophore. 11. Columella on long sporangiophore. 12. Sporangio-
spores. 1, 9 bars = 20 um; 2-8, 10-12 bars = 10 um.
Mucor circinelloides f. janssenii (Lendn.) Schipper, Stud. Mycol. 12: 13. 1976.
PLATE 2 (FIGS. 9-12)
SPECIMEN EXAMINED: Brazil, Sao Paulo State, Santa Gertrudes, urban area, 22°27'12”S
47°32’39"W, soil from a public garden, j.. de Souza & N.C. Schoenlein, 18.11.2003, No.
2036.
Colonies 5 days old on SMA at 25 °C, obverse powdery graphite grey with a
pale beige to white margin, or powdery light orange grey without a margin,
reverse pale beige, light grey or orange, 60 x 55 mm in size, up to 3 mm high;
sporangiophores sympodially branched, slightly curved to circinate, with
reduced apical width (short) or straight (long), septate or non-septate, 15-
267(-821) wm long, 4.5-10 um in diameter at base and 3-8.7 um in diameter
at apex; sporangia globose, slightly applanate, sporangial wall deliquescent,
pale chestnut, dark chestnut to almost black, 16.5-56 x 18-57 um; columellae
globose, slightly applanate to ellipsoidal, collars present, with or without
granular contents, hyaline to slightly grey, may be somewhat light greenish,
11.2-35 x 17-35 um; sporangiospores globose, ovoid to almost spherical,
Mucorales from contaminated soils (Brazil) ... 279
smooth-walled, chestnut in mass, pale chestnut when individual, 3-6.7 x
2.2-6 um; chlamydospores apical or intercalate, 9.7-23.2 x 8.2-16.5 um;
submerged mycelium with olive green contents, oidia present; zygosporangia
not observed.
Notes. Isolated on MA with 100 mg of Pb(NO,),. After 5 days on SMA at 20
and 30 °C growth and sporulation; after 5 days at 37 °C very poor growth and
no sporulation but the culture grew when placed again at 25 °C. This form of
M. circinelloides was isolated from soil (Schoenlein-Crusius et al. 1996, 2006),
water, soil and leaves (Schoenlein-Crusius & Milanez 1997) and herbivore dung
(Alves et al. 2002, Santiago 2008).
Mucor hiemalis Wehmer, Annls. Mycol. 1: 37. 1903.
f. hiemalis PLATE 3 (FIGS. 1-4)
SPECIMENS EXAMINED: Brazil, Sao Paulo State, Santa Gertrudes, urban area, 22°27'12”S
47°32°39°W, soil from a public garden; Fazenda Itaqui drainage basin, contaminated
area near ceramic tile industries, 22°27°49"S 47°32’32”W, soil under grass, J.J. de Souza
e N.C. Schoenlein, 28.V.2003, No. 2037-2039; contaminated area near ceramic tile
industries, 22°27'46"S 47°32’29"W, 22°27'58°S 47°32’22”W, soil under grass; J.I. de
Souza & N.C. Schoenlein, 30.V1.2003, No. 2040, No. 2041.
Colonies 5-7 days old on SMA at 25 °C, obverse powdery, pale beige dotted
in grey, or pale orange slightly white, reverse pale beige to pale orange with
or without a deep orange (peach) coloration on inoculum site, 90 mm in
diameter, 5-14 mm high; sporangiophores sympodially branched, straight
to slightly curved, arising from the main branches, branches septate or non-
septate, with or without granular orange contents, 34-2,650 um long, 4-18.7
[Lm in diameter at base and 2.3-16.2 um in diameter at apex; sporangia globose,
slightly applanate, with sporangial wall highly deliquescent at 5 days, golden
yellow, chestnut (amber) to greenish chestnut, 29-89 x 30-88 um; columellae
ellipsoidal, base may be truncate, pyriform, or globose, slightly applanate,
collars present or absent, hyaline green to slightly grey, with or without granular
contents, 7.2-56 x 7.5-46 um; sporangiospores ellipsoidal, smooth, flattened at
one side, with homogeneous or granular contents, pale green, with or without
dark green droplets, 3-10.9 x 1.8-6.8 um; chlamydospores apical, catenulate
or intercalate, present or absent on the aerial mycelium, with or without olive
green granular contents, 6.2-30 x 5-24.4 um; submerged mycelium with olive
green to greenish golden yellow contents, with or without rhizoid-like thin side
branches, oidia present or absent; zygosporangia not observed.
Notes. Isolated on MA metal free, with 100 mg of Pb(NO,),, and 200 mg of
Zn(NO,),. No strain grew after 7 days on SMA at 37 °C. The form hiemalis was
isolated from soil (Schoenlein-Crusius et al. 1996, Santos et al. 1998, Ruegger
et al. 2002, Tauk-Tornisielo et al. 2005), water, soil and leaves (Schoenlein-
280 ... de Souza & al.
Crusius & Milanez 1997), soil and leaves (Schoenlein-Crusius et al. 2006), and
herbivore dung (Alves et al. 2002, Santiago 2008). M. hiemalis was isolated from
soil (Trufem 1978, 1981la, Antunes et al. 1993), leaves (Schoenlein-Crusius &
Milanez 1989, Wellbaum et al. 1999) and herbivore dung (Trufem 1984, Trufem
& Viriato 1985).
Mucor hiemalis f. luteus (Linnem.) Schipper, Stud. Mycol. 4: 33. 1973.
PLATE 3 (FIGS. 5-8)
SPECIMENS EXAMINED: Brazil, Sao Paulo State, Santa Gertrudes, Fazenda Itaqui drainage
basin, contaminated area near ceramic tile industries, 22°27'46”S 47°32’29” W, soil under
grass, J.J. de Souza & N.C. Schoenlein, 28.V.2003, No. 2042-2046, 30.VI.2003, No. 2047.
Colonies 5-7 days old on SMA at 25 °C, sometimes with a strong baker’s yeast
odor, obverse pale beige to slightly orange to pale whitish orange, or containing
a 50 mm central region up to 14 mm high of cotton-like mycelium greyish
orange dotted in graphite grey, with a 8 mm high peripheral region with or
without sectors of whitish dense mycelium, reverse orange, deep orange (peach)
or yellow mango, 90 mm in diameter, up to 14 mm high; sporangiophores .
sympodially branched, septate or non-septate, sometimes curved, 25-5,750
um long, 5.6-18.4 (—22.5) um in diameter at base and 3.5-13.8 (-20) um in
diameter at apex; sporangia globose, slightly applanate, sporangial wall highly
deliquescent, amber to greyish-greenish chestnut, 23.7-80 x 22.5-85 um;
columellae globose, slightly applanate, spherical or ellipsoidal, collars present
or absent, with or without granular contents, hyaline green (minor) to slightly
grey (major), 12.5-66 x 11.3-68 um; sporangiospores narrow ellipsoidal,
sometimes flattened at one side, some slightly curved, smooth-walled, with or
without granular contents, pale green, 3.2-13.6 x 1.7-6.4 um; chlamydospores
catenulate, isolated, apical or intercalate, present or absent on aerial mycelium,
7.2-30 x 6.2-21.7 um; submerged mycelium with olive green to golden yellow
contents, sometimes with thin side branches, oidia absent; zygosporangia not
observed.
Notes. Isolated on MA with 100, 200 and 500 mg of Zn(NO,),. No strain grew
after 7 days on SMA at 37 °C; the strain No. 2044 showed reduced growth but
no sporulation. This form of M. hiemalis was isolated from soil (Schoenlein-
Crusius et al. 1996), air (Schoenlein-Crusius et al. 2001), soil and leaves
(Schoenlein-Crusius et al. 2006) and herbivore dung (Alves et al. 2002).
Mucor racemosus Fresen., Beitr. Mykol. 1: 12. 1850.
f. racemosus PLATE 3 (FIGS. 9-12)
SPECIMEN EXAMINED: Brazil, Sao Paulo State, Santa Gertrudes, Fazenda Itaqui drainage
basin, contaminated area near ceramic tile industries, 22°27'46”S 47°32’29” W, soil under
grass, J.I. de Souza & N.C. Schoenlein, 30.V1.2003, No. 2048.
Mucorales from contaminated soils (Brazil) ... 281
@
PLATE 3. FIGS. 1-4. Mucor hiemalis f. hiemalis: 1. Sporangium. 2, 3. Columellae. 4. Sporangiospores.
FIGs. 5-8. M. hiemalis f. luteus: 5. Sporangium. 6, 7. Columellae. 8. Sporangiospores. FIGS. 9-12.
M. racemosus f. racemosus: 9, 10. Short sympodial branches. 11. Columella. 12. Sporangiospores.
FIGS. 13-16. Zygorhynchus moelleri: 13. Sporangium. 14. Zygosporangia. 15. Columella.
16. Sporangiospores. 1-9, 11-13, 15, 16 bars = 10 um; 10, 14 bars = 20 um.
Colonies 5 days old on SMA at 25 °C, scarce mycelium, obverse pale greenish
grey, powdery, dotted in graphite grey, some colonies with more dense and
higher (7 mm) orange mycelium sectors, reverse grey slightly orange, 90 mm
in diameter, 5 mm high; sporangiophores sympodially branched, short or long,
septate or non-septate, 35-1,020 um long, 7.5-13.8 um in diameter at base and
6.3-11.3 um in diameter at apex; sporangia globose, applanate, sporangial wall
persistent or deliquescent from sporangia borned on short or long branches,
respectively, pale chestnut (amber), dark chestnut to almost black, 28-90 x 30-
95 um; columellae ellipsoidal or globose, applanate, mostly with collars, hyaline
green to slightly grey, 15-51 x 15-43 «tm; sporangiospores wide ellipsoidal,
pale green to slightly chestnut, 4.5-10.9 x 3.6-7.7 um; chlamydospores isolated,
282 ... de Souza & al.
catenulate or apical, 6.2-25 x 7.5-18.7 um; submerged mycelium with olive
green contents and terminating in oidia; zygosporangia not observed.
Notes. Isolated on MA with 100 mg of Pb(NO,),. After 7 days on SMA at 37
°C no growth, but both growth and sporulation started again when the cultures
were placed at 25 °C. The form racemosus was isolated from soil (Trufem 1978,
Schoenlein-Crusius et al. 1996), herbivore dung (Viriato 2003) and soil and
leaves (Schoenlein-Crusius et al. 2006). M. racemosus was isolated from soil
(Upadhyay 1967, Santos et al. 1989, Antunes et al. 1993).
Zygorhynchus moelleri Vuill., Bull. Soc. Mycol. France 19: 117. 1903.
PLATE 3 (FIGS. 13-16)
SPECIMEN EXAMINED: Brazil, Sao Paulo State, Santa Gertrudes, Fazenda Itaqui drainage
basin, contaminated area near ceramic tile industries, 22°27°49"S 47°32’32” W, soil under
grass, J.I. de Souza & N.C. Schoenlein, 18.11.2003, No. 2051.
Colonies 5 days old on SMA at 25 °C, obverse light greyish white, varying
from pale grey sand to graphite grey in 25-30 days, highly dotted in black, |
dry mycelium, reverse pale greenish beige, 90 mm in diameter, 9 mm high,
up to 12 mm high in 25-30 days; sporangiophores sympodially branched,
slightly curved, some very short bearing almost sessile sporangia, 31-250 um
long, 3.7-12.5 um in diameter at base and 2.5-12.5 um in diameter at apex;
sporangia globose, pale chestnut brilliant (amber), sporangial wall persistent,
smooth, 15-48 x 15-46 um; columellae applanate, greenish hyaline to slightly
chestnut, 9-30 x 19.5-44 um; sporangiospores ellipsoidal, some curved, to
bean-shaped, pale green, mostly showing olive green droplets in the apices,
3.7-6.7(-11.2) x 2.2-3.7 um; chlamydospores rare; submerged mycelium
with olive green brilliant contents; zygosporangia very abundant, globose to
slightly applanate, pale chestnut (amber) when young, dark chestnut to black
when mature, 30-45 x 32-47 um, with spines 5-7(-10) um high; suspensors
unequal, one thin, long, straight and other inflated and bending to the former;
homothallic; azygosporangia not observed.
Notes. Isolated on MA with 100 mg of Pb(NO.,).. Brazilian isolates of Z. moelleri
from soil were studied by Hesseltine et al. (1959). Other authors isolated it from
soil (Trufem 1978, Antunes et al. 1993, Schoenlein-Crusius et al. 2006), leaves
(Schoenlein-Crusius et al. 1990) and water, soil and leaves (Schoenlein-Crusius
& Milanez 1997).
Absidia cylindrospora Hagem, Skr. Vidensk.-Selsk. Christiania, Kl. I, Math.-Natur.
1907(7): 45. 1908. var. cylindrospora PLaTE 4 (FIGs. 1-4)
SPECIMENS EXAMINED: Brazil, Sao Paulo State, Santa Gertrudes, Fazenda Itaqui drainage
basin, contaminated area near ceramic tile industries, 22°27°59"S 47°32'22”W, soil under
-
LN ee
Mucorales from contaminated soils (Brazil) ... 283
PLATE 4. FIGS. 1-4. Absidia cylindrospora v. cylindrospora: 1. Verticil. 2. Sporangium. 3. Columella.
4. Sporangiospores. FIGS. 5-8. Rhizopus arrhizus v. arrhizus: 5. Sporangium. 6, 7. Columellae.
8. Sporangiospores. FIGS. 9-12. Cunninghamella phaeospora v. phaeospora: 9. Sporangiophore
monopodially branched. 10. Terminal vesicle and lateral branches. 11, 12. Sporangiola unispored.
1, 3, 5-7 bars = 20 um; 2, 4, 8, 9-12 bars = 10 um.
grass, J.I. de Souza & N.C. Schoenlein, 18.III.2003, No. 2031; urban area, 22°27'12°S
47°32°39’W, soil from a public garden, J.I. de Souza & N.C. Schoenlein, 18.1II.2003, No.
2030.
Colonies 5 days old on SMA at 25 °C, obverse greyish white, chestnut in 25-
30 days, reverse wavy zonate, pale creamy beige, 76 mm in diameter and 15
mm high; sporangiophores pale chestnut, pale beige, or hyaline green when
small, arising singly, in pairs or in verticils 3 or 4(-5), produced on stolons,
50-364 um long, 2.5-8.2 ttm in diameter at base and 2.2—5.5 um in diameter at
apex, septate; sporangia spherical at the apices, conical at the base, apophysate,
smooth, pale olive green to chestnut, sporangial wall deliquescent, 12.5-30
x 13.8-33 um; columellae rounded, with or without 1.2-6.2 um thin erect
projections sometimes inflated at the apex, sporangial membranes between the
columella-apophysis region may persist, pale olive green to pale beige, 6.2-20
x 10-22.5 um; sporangiospores cylindrical with round extremities, pale green,
3.5-5.7 x 2.2-3.2 um; chlamydospores apical or isolated, scarce, 4.5-17.5 x 4.5-
284 ... de Souza & al.
10.5 um; submerged mycelium containing olive green droplets; zygosporangia
not observed.
Notes. Isolated on MA with 100 mg of Pb(NO,),, and 200 mg of Zn(NO,)..
After 5 days on SMA at 30 °C slightly decreased growth and normal sporulation;
after 7 days at 37 °C no growth; after 5 days at 25 °C on malt extract agar
pale brownish-greenish grey colonies were developed, indicating the variety
cylindrospora (Schipper 1990). Brazilian isolates of the variety cylindrospora
were obtained from Brazil nuts, soil (Hesseltine & Ellis 1964) and soil (Pfenning
1993). Isolates of A. cylindrospora were obtained from soil (Upadhyay 1970,
Lira 1971, Trufem 1978, 1981b, Antunes et al. 1993, Schoenlein-Crusius et al.
1996, Santiago & Souza-Motta 2006), soil and leaves (Schoenlein-Crusius &
Milanez 1997), soil and herbivore dung (Viriato 2003) and corn flour (Ribeiro
et al. 2003).
Rhizopus arrhizus A. Fisch., Rabenh. Krypt.-Fl. Ed. 2, 1(4): 233. 1892.
var. arrhizus PLATE 4 (FIGS. 5-8) _
= Rhizopus oryzae Went & Prins. Geerl., Verh. K. Nederl. Akad. Wet., 2 Sect. 4(2): 16. 1895.
SPECIMENS EXAMINED: Brazil, Sao Paulo State, Santa Gertrudes, Fazenda Itaqui drainage
basin, contaminated area near ceramic tile industries, 22° 27’ 46” S 47° 32’ 29” W, soil
under grass, J.J. de Souza & N.C. Schoenlein, 30.V1.2003, No. 2049, No. 2050.
Colonies 7 days old on SMA at 25 °C, obverse greyish white densely dotted with
black sporangia, reverse greyish creamy white, sometimes yellowish, 90 mm
in diameter, 15 mm high; sporangiophores arising from stolons connected to
substrate by rhizoids, sometimes almost verticillate, arising from vesicle apices
of wide hyphae, or on isolated dichotomous branches, chestnut to hyaline
green near substrate, some entirely hyaline green, 370-1,600 um long, 7.5-20
ttm in diameter at base and 15-30 um in diameter at apex; sporangia globose,
apophysate, slightly olive green when young, grey to black when mature,
sporangial wall persistent and smooth, 50-160 x 60-165 um; columellae
ellipsoidal with truncate base or globose, without collars or with great remaining
sporangial membranes adhered to the base, pale chestnut to hyaline green,
27.5-120 x 37.5-120 um; sporangiospores ellipsoidal/subglobose angular, thick
walled, slightly striate, greenish to slightly chestnut when solitary, greenish grey
when in mass, sometimes in pairs, 5.5-12 x 4-9 um; chlamydospores absent;
submerged mycelium with rhizoids; zygosporangia not observed.
Notes. Isolated on MA with 200 mg of Pb(NO,),. After 7 days on SMA at 45 °C
poor growth and no sporulation; after 7 days at 37 °C slightly decreased growth
and sporangia without free sporangiospores. R. arrhizus was isolated from soil
(Trufem 1981b), water, soil and leaves (Schoenlein-Crusius & Milanez 1997)
and herbivore dung (as R. oryzae, Santiago 2008).
Mucorales from contaminated soils (Brazil) ... 285
Cunninghamella phaeospora Boedijn, Sydowia 12: 348. 1959 [‘1958’].
var. phaeospora PLATE 4 (FIGS. 9-12)
SPECIMEN EXAMINED: Brazil, Sao Paulo State, Santa Gertrudes, urban area, 22°27°12”S
47°32°39°W, soil from a public garden, J.J. de Souza & N.C. Schoenlein, 18.11.2003, No.
2032.
Colonies 5 days old on SMA at 25 °C, like moist cotton, obverse greyish white,
to pale grey in 25-30 days, reverse wavy zonate, creamy white, to greyish creamy
beige in 25-30 days, 84 x 61 mm in size and 16 mm high; aerial mycelium
abundant, 6.2-22.5 um in diameter; sporangiophores hyaline to olive green,
septate or non-septate, monopodially branched (very long), with isolated
dichotomous branches, lateral branches, or arising from stolons (minority),
rarely pseudoverticillate, up to 90 um long, 7.5-13.7 um in diameter, sympodial
branches absent; terminal vesicles hyaline to olive green, 17.5—46 tum in diameter
at the wider part, lateral small vesicles on aerial mycelium up to 11.2 um in
diameter; pedicels up to 5 um long; sporangiola unispored, almost spherical
or globose, rarely lemon-shaped, hyaline green to olive green or pale chestnut,
12.5-18.8 x 12.2-18.2 um, mostly spineless, or rarely with spines 1.8-3.6 um
long; chlamydospores absent; rhizoids sometimes finger-like; submerged
mycelium with inflated regions sometimes with olive green drops, ending with
thin hair-like irregular branches; zygosporangia not observed.
Notes. Isolated on MA with 100 mg of Zn(NO,),. After 5 days on SMA at
20 and 30 °C normal growth and sporulation; after 7 days at 37 and 45 °C no
growth. The strain No. 2032 generally formed globose sporangiola (12.5-18.8
x 12.2-18.2 um), which are relatively larger than the globose, lacrymoid and
ovoid (6.6-16.0 um) ones that has been observed (Baijal & Mehrotra 1980,
Lunn & Shipton 1983, Zheng & Chen 2001). C. phaeospora was isolated from
soil (Upadhyay 1970, Santos et al. 1998, Viriato 2003, Maia et al. 2006).
Acknowledgements
The authors are very grateful to Dr Gerald L Benny (University of Florida) and Dr Kerry
O'Donnell (United States Department of Agriculture) for the critical reviewing of the
manuscript and valuable suggestions, Roberto J. Pedro (Universidade Estadual Paulista)
for technical support on soil collections, Dr Ronaldo NM Pitombo and Gledson M
Guimaraes (Universidade de Sao Paulo) for freeze-drying the fungal cultures, Dr Marcelo
P Marcelli and his team (Instituto de Botanica) for skilled help with the manuscript and
preparation of the figures, and to CAPES (Coordenacao de Aperfeigoamento de Pessoal
de Nivel Superior) for the doctoral scholarship provided to JI de Souza.
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Wellbaum C, Schoenlein-Crusius IH, Santos VB. 1999. Fungos filamentosos em folhas do ambiente
terrestre e aquatico da Ilha dos Eucaliptos, Represa do Guarapiranga, Sao Paulo, SP. Revista
Brasileira de Botanica 22: 69-74.
Zheng RY, Chen GQ. 2001. A monograph of Cunninghamella. Mycotaxon 80: 1-75.
MYCOTAXON
Volume 106, pp. 289-295 October-December 2008
Validation and typification of Laetiporus montanus
MICHAL TOMSOVSKY & LIBOR JANKOVSKY
tomsovsk@mendelu.cz
Faculty of Forestry and Wood Technology
Mendel University of Agriculture and Forestry in Brno
Zemeédelska 3, CZ-613 00, Brno, Czech Republic
Abstract — Laetiporus montanus (Basidiomycota, Polyporales) is an invalidly described
taxon growing on conifers in Central European mountain areas. A molecular
phylogenetic study of the species was performed to reveal its taxonomical position. The
sequencing of ITS and LSU regions of ribosomal DNA revealed high proximity of the
sequences of L. montanus to the North American species L. huroniensis. Nevertheless,
DNA sequences of the two taxa are not identical and these species also differ in
basidiospore dimensions. The ITS sequence of L. montanus is also identical to Japanese
specimens named as L. sulphureus var. miniatus. However, the basionym of this taxon,
Polyporus miniatus, was described from the tropics and its identity with temperate
conifericolous specimens is improbable. Therefore, Laetiporus montanus sp. nov. is
validated and described here.
Key words — Basidiomycetes, Picea, Larix, taxonomy, polyporoid clade
Introduction
Laetiporus sulphureus s.1. is a large, yellow to orange polypore, considered to be
widely distributed from boreal to tropical zones. It grows on different broadleaved
and coniferous hosts causing brown rot of wood, occasionally resulting in
economical losses. As in many other fungal species, Laetiporus sulphureus
s.l. is formed of a complex of several sibling species. North American studies
revealed the presence of five species and one variety within the L. sulphureus
complex (Burdsall & Banik 2001). In Central Europe a morphologically and
ecologically distinguishable species, Laetiporus montanus, was described by
Cerny (1989). However, the name Laetiporus montanus Cerny was published
invalidly, because a Latin diagnosis and an indication of the type are lacking
in the publication (Cerny 1989). Nevertheless, the taxon seems to be a good
species restricted to coniferous wood in mountain areas, differing from other
Central European Laetiporus specimens by its larger basidiospores. The aim
of our work is to confirm the taxonomic position of L. montanus according to
sequences of ribosomal DNA.
290 ... Tomsovsky & Jankovsky
Material and methods
To assure the taxonomic position of Laetiporus montanus, sequencing of the ITS and
LSU regions of nuclear ribosomal DNA was carried out.
A representative specimen of L. montanus, collected in the locality mentioned in
original publication (Cerny 1989), deposited in BRNM (no. 706688), and the respective
culture deposited in MUAF culture collection (Collection of Mendel University of
Agriculture and Forestry in Brno, Department of Forest Protection and Wildlife
management), isolate no. 856, were selected for the study. The specimen was sequenced
together with a sample of L. sulphureus s.s., collected in the Czech Republic on Sorbus
aucuparia, deposited in BRNM (specimen 706696).
DNA was isolated from fresh culture and from dried tissue of herbarium specimens
using PowerSoil™ DNA Isolation Kit (Mo-Bio). The DNA was amplified with PCR,
using the Mastercycler® ep thermocycler (Eppendorf). The PCR amplifications were
performed according to TomSovsky et al. (2006).
Sequences were determined with an ABI PRISM 3100 Avant DNA sequencer
(Applied Biosystems) at the Department of Animal Morphology, Physiology and
Genetics, Faculty of Agriculture, Mendel University, Brno using the ABI PRISM BigDye
terminator v1.1 cycle sequencing kit (Applied Biosystems). All samples were sequenced ~
with the primers used in the PCR.
GenBank search of newly obtained ITS and LSU sequence of L. montanus using
BLAST (Altschul et al. 1990) resulted in 99-100 % identity with those of the North
American conifer-inhabiting species Laetiporus conifericola Burds. & Banik and
Laetiporus huroniensis Burds. & Banik (GenBank Access. Nos. EU402523-24,
EU402539-40, EU402569-78; Lindner & Banik 2008). In the case of L. montanus
ITS sequences, 100 % identity with Japanese specimens of L. sulphureus var. miniatus
(GenBank Accession Nos. AB308192-95; Ota & Hattori 2008) was revealed.
Newly obtained sequences were edited manually using BioEdit version 4.7.1.
(Hall 1999), and selected L. conifericola, L. huroniensis and L. sulphureus var. miniatus
sequences from the GenBank (GenBank Accession Nos. see at Fig. 1) were added to the
dataset. The combined ITS and LSU alignment with introduced gaps consisted of 1183
characters including 1163 constant and 20 variable sites.
A neighbor-joining phylogenetic analysis was carried out using MEGA 4 (Tamura
et al. 2007). The phylogram was constructed using the maximum composite likelihood
model and the support for the topology of the dendrogram was estimated using 1000
bootstrap-replicates.
Results and discussion
The newly obtained Laetiporus montanus sequences of both gene regions show
high similarity to the North American species L. huroniensis, while L. conifericola
forms a sister clade (Fic. 1). However, there are some differences between
the sequences of L. montanus and L. huroniensis (TABLES 1, 2) and also their
basidiospore dimensions differ. L. montanus has broadly ovoid basidiospores,
(5.8) 6.0-8.0 (9.0) x (3.8) 4.0-5.5 um (Cerny 1989; own observations), while
those of L. huroniensis are smaller (5.0-7.0 x 4.2-5.0 um; according to Burdsall
Laetiporus montanus validated ... 291
L.. montanus EU840558/EU884419
L. sulphureus var. miniatus
AB308192/-
L. huroniensis EU402573/EU402539
L. huroniensis EU402571/EU402540
L. conifericola
EU402577/EU462524
L. conifericola
EU402575/EU402523
L. sulphureus
EU840565/EU884420
0.001
FiGure 1. Phylogenetic tree of Laetiporus montanus and related taxa based on combined sequences
of the ITS and LSU regions of nuclear ribosomal DNA region (the first GenBank accession number
refers to the ITS sequence the second one to LSU sequence). The tree was constructed by a
neighbor-joining analysis; bootstrap values are shown at the nodes. The bar indicates the number
of substitutions per position.
TABLE 1. Comparison of nucleotide polymorphism of aligned conifericolous
Laetiporus ITS sequences.
1-5 15 413 507-513
TTCGGAC
L. sulphureus var. ACGAA C T G TTCGGAC
miniatus AB308192
| : huroniensis
: U402571
L. conifericola T C A TTCGGAC
402577
.. conifericola T C A TTEEGAC
EU402575
TABLE 2. Comparison of nucleotide polymorphism of aligned conifericolous
L. huroniensis RCO C 1 G
EU402573
Laetiporus LSU sequences.
L. montanus GGATTCC
EU884419
L. huroniensis GGATTCC T
EU402539
L. huroniensis GGATTCC i
EU402540
S
eagGe
een a ee
es Hina tio et
ied i
eve
L. conifericola GGATTCC
EU402524
L. conifericola GGATTCC
EU402523
=
292 ... TomSovsky & Jankovsky
& Banik 2001). Basidiospore dimensions of L. conifericola are almost identical
to those of L. montanus (6.5-8.0 x 4.0-5.0 um), but the two species differ in
their DNA sequences more than L. huroniensis differs from L. montanus.
The ITS sequences of L. montanus and Japanese material of L. sulphureus
var. miniatus are identical. In addition, colour of basidiocarp surface and
pores, and substrate specificity of these two taxa are almost identical (Ota &
Hattori 2008; Cerny 1989). Unfortunately, Ota & Hattori’s publication lacks
basidiospore dimensions and LSU sequence data, which could be used to
assure the identity of Central European L. montanus and these Japanese fungi.
The name L. sulphureus var. miniatus (Jungh.) Imazeki is currently used for
various morphologically and ecologically different specimens occuring mainly
in temperate or boreal Japan (Ota & Hattori 2008). The basionym of this taxon,
Polyporus miniatus Jungh., was described from tropical Asia (island of Java).
Moreover, Corner’s specimens of L. sulphureus var. miniatus from Papua
and the Solomon Islands (Corner 1984) are characterized by basidiospores
significantly narrower [4-5.5(-5.7) x 3-3.7 um] than those of the European
taxa. Therefore the conspecificity of the Japanese temperate—boreal Laetiporus
specimens growing on coniferous trees and L. sulphureus var. miniatus sensu
typi is improbable. According to the above evidence the re-description and
typification of Laetiporus montanus is proposed.
Laetiporus montanus Cerny ex TomSovsky & Jankovsky, sp. nov. FIGURE 2
MycoBank : MB 512119
Species haec a Laetiporo sulphureo basidosporis late ovoideis, (5.8) 6.0-8.0 (9.0) x (3.8)
4.0-5.5 um differt. Habitat ad substrato arboribus coniferis (Picea, Larix) in partibus
montanis Europae Centralis.
Ho.ortypus: Czech Republic, Jeseniky Mts., Pradéd national nature reserve, valley of
the Bila Opava brook, 50°4’28.35"N; 17°14’41.17”E, on stump of Picea abies, 1230 m
a.s.L, 27 July 2006, leg. L. Jankovsky, BRNM 706688.
ETyMoLoey: from the Latin montanus = occurring in mountains.
PREVIOUSLY PUBLISHED DESCRIPTIONS AND ILLUSTRATIONS: Cerny (1989, Parazitické
drevokazné houby: 45-47).
KEY CHARACTERS — Basidiocarps annual, shelf-shaped, dimidiate, sessile
to stipitate, up to 30 cm wide, projecting 3-20 cm and 1-3 cm thick. Upper
surface of basidiocarp bright orange, fading to pale brownish with age; context
pale yellow; pore surface bright sulphureous yellow to pale tan with age; pores
angular, 1-4 per mm, with thin dissepiments that become lacerate; tube layer
2-4 mm thick. Hyphal system dimitic; generative hyphae rarely apparent in
the context, thin-walled, hyaline, simple-septate with rare branching, 4-8 um
in diam. Contextual binding hyphae firm to thick walled, hyaline, nonseptate,
much branched, 5-13 um in diam; tramal hyphae thin to thick walled,
simple-septate with occasional branching, 3-6 um in diam. Basidia pyriform,
Laetiporus montanus validated ... 293
B
(0!
A C
5m
FiGuRE 2. Microscopic characters of Laetiporus montanus.
A - thin-walled generative hypha; B - contextual binding hypha; C - basidiospore; D - basidium.
4-sterigmatic, 12-25 x 6-8 um. Basidiospores broadly ovoid, (5.8) 6.0-8.0 (9.0)
x (3.8) 4.0-5.5 um, hyaline, thin-walled, smooth, negative in Melzer’s reagent,
usually with globose oil drop, 1-3 um in diam. For microscopic characters,
pee FIG.2.
ECOLOGY, DISTRIBUTION — Laetiporus montanus occurs on mature and
Overmature conifers (Picea abies and Larix decidua) in mountain areas of
Central Europe. The species seems to be restricted to the elevation above 1100
m a. s. |. and to stands of indigenous coniferous trees. The currently known
distribution is restricted to two localities in the Czech Republic: Jeseniky Mts.
(Altvatergebirge in German) and Bohemian Forest (Sumava Mts. in Czech;
Bohmerwald in German), to High Tatra Mts. in Slovakia, and large areas in the
Austrian Alps. In the Czech Republic, the occurrence L. montanus is strictly
restricted to autochthonous montane spruce stands and natural range of Picea
abies.
REPRESENTATIVE SPECIMENS EXAMINED — Laetiporus montanus: CZECH REPUBLIC:
Jeseniky Mts., Pradéd national nature reserve, 1320 m a.s.l., Picea abies, 27.V1I.2006,
leg. L. Jankovsky, holotype, BRNM 706688, isotype BRNL. - Bohemian Forest (Sumava
Mts.), Plechy Mt., 1340 m a.s.l., Picea abies, 15.VII.1998, leg. J. Holec, PRM 897053. -
Bohemian Forest (Sumava Mts.), Trojmezna Mt., 1270 ma.s.l., Picea abies, 13.V1I.2006,
leg. V. Pouska, BRNM 706697. SLOVAKIA: High Tatra Mts., Popradské Pleso Lake,
ca.1400 m a.s.l., Larix decidua, 30.V1I.1987, leg. A. Cerny, BRNL. AUSTRIA: Upper
Austria, Burgtall, Wurzeralm, 1700 ma.s.l., Larix decidua, 17.1X.1989, leg. H. Forstinger,
LI. - Styria, surroundings of Oberw6lz, 2000 m a.s.l., Larix decidua, 30.V.1971, leg. H.
Forstinger, LI.
294 ... TomSovsky & Jankovsky
Laetiporus sulphureus: CLECH REPUBLIC: Tisnov town, Prunus domestica, 3.X.2006,
leg. L. Jankovsky, BRNM 706687. - Ceskosaské Svycarsko National Park, Mezni{ louka,
host not indicated, 3. VII.2003., leg. A. Vagner, BRNM 686283. - Orlické Mts., Bukacka
protected nature area, 1000 maz.s.l., Sorbus aucuparia, 1.1X.2006, leg. J. Moravec, BRNM
706696. - Beskydy Mts., Komorni Lhotka village, 415 m a.s.l., Larix decidua, 3.V1.2006,
leg. J. Lederer, BRNM 706686.
COMMENTS - The distribution of Laetiporus montanus is poorly known. The
species occurs probably not only in Austria but also in Switzerland (Rogers et
al. 1999) and other adjacent Alpine countries. Its presence in other European
mountain areas with indigenous Picea and Larix is possible. In contrast,
misidentifications of L. sulphureus specimens fruiting on planted conifers at
lower elevations are possible (e.g. BRNM 706686), even though the two species
are distinguishable according to basidiospore measurements - the length of
L. sulphureus basidiospores usually does not exceed 7.0 um.
The occurrence of L. montanus in Asia, mainly in Siberia, the Russian Far
East and northern Korea or Japan is possible. There may be rich populations of
this or a closely related species occurring in east Palaearctic regions with natural
stands of coniferous trees. Therefore a study of conifericolous Laetiporus from
these regions is desirable.
Acknowledgments
The work was supported by the Ministry of Education, Youth and Sports of the Czech
Republic, project no. MSM 6215648902. We are grateful to Dr. Vladimir Antonin
(Moravian Museum, Brno, the Czech Republic) and Dr. Tuomo Niemela (Finnish
Museum of Natural History, University of Helsinki, Finland) for presubmittal review
of the manuscript.
Literature cited
Altschul SF, Gish W, Miller W, Myers EW, Lipman DJ. 1990. Basic local alignment search tool. J.
Mol. Biol. 215: 403-410.
Burdsall HH, Banik MT. 2001. The genus Laetiporus in North America. Harvard Papers in Botany
6: 43-55.
Cerny A. 1989. Parazitické dievokazné houby. Statni zemédélské nakladatelstvi: Praha. 104 pp.
Corner EJH. 1984. Ad Polyporaceas II & III. Nova Hedwigia Beihefte 78: 1-222.
Hall TA. 1999. BioEdit: a user-friendly biological sequence alignment editor and analysis program
for Windows 95/98/NT. Nucleic Acids Symp. Ser. 41: 95-98.
Lindner DL, Banik MT. 2008. Molecular phylogeny of Laetiporus and other brown-rot polynom
genera in North America. Mycologia 100: 417-430.
Ota Y, Hattori T. 2008. Relationships among three Japanese Laetiporus taxa based on phylogenetic
analysis and incompatibility tests. Mycoscience 49: 168-177.
Rogers SO, Holdenrieder O, Sieber T. 1999. Intraspecific comparisons of Laetiporus sulphureus
isolates from broadleaf and coniferous trees in Europe. Mycological Research 103:1245-1251.
Ryvarden L, Gilbertson RL. 1993. European polypores, Part 1. Fungiflora: Oslo: 393 pp.
Laetiporus montanus validated ... 295
Tamura K, Dudley J, Nei M, Kumar S. 2007. MEGA4: Molecular Evolutionary Genetics Analysis
(MEGA) software version 4.0. Molecular Biology and Evolution 24: 1596-1599.
Tomsovsky M, Kolatik M, Pazoutova S, Homolka L. 2006. Molecular phylogeny of European
Trametes (Basidiomycetes, Polyporales) species based on LSU and ITS (nrDNA) sequences.
Nova Hedwigia 82: 269-280.
MY COTA XON
Volume 106, pp. 297-302 October-December 2008
Two little known gasteroid fungi
from Santa Catarina State, southern Brazil
VAGNER GULARTE CORTEZ’, MARCELO ALOISIO SULZBACHER?’,
IURI GOULART BASEIA? & ROSA MARA BORGES DA SILVEIRA!
cortezvg@yahoo.com.br; baseia@cb.ufrn.br
‘Universidade Federal do Rio Grande do Sul, Departamento de Botanica
Av. Bento Goncalves 9500, 91501-970, Porto Alegre, RS, Brazil
Rua Garibaldi 7, 96815-700, Santa Cruz do Sul, RS, Brazil
3Universidade Federal do Rio Grande do Norte
Departamento de Botanica, Ecologia e Zoologia
59072-970, Natal, RN, Brazil
Abstract — Geastrum ovalisporum and Tulostoma rickii are reported for the first
time from the State of Santa Catarina in southern Brazil. Geastrum ovalisporum is
characterized mainly by ovoid basidiospores, a unique feature in the genus. Tulostoma
rickii, a rare and poorly known species from Rio Grande do Sul State, is diagnosed
by its bulbous (pseudo-volvate) stipe base and reticulate basidiospores. Illustrations
(including scanning electron micrographs of the basidiospores) accompany full
taxonomic descriptions and distributional notes.
Key words — Geastraceae, Tulostomataceae, Neotropical mycobiota
Introduction
The study of the gasteroid mycobiota of Santa Catarina State dates from the
/ mycological contributions of the European naturalists who explored Brazil
| in the past centuries, such as E. Ule and A. Mdller (Fidalgo 1965). The latter
| studied species of phalloids, including the new genera Blumenavia, Itajahya,
-and Protubera, which were based on specimens collected in Santa Catarina
| (Moller 1895). Viégas (1945) reported two unidentified species of Cyathus and
Lycoperdon for the State. More recently, Giachini et al. (2000) surveyed the
ectomycorrhizal fungi associated to exotic Pinus and Eucalyptus, reporting 23
| gasteroid fungi, including three new species, and Sobestiansky (2005) reported
collections of Geastrum triplex Jungh. and Sphaerobolus sp. The discussion
| of two noteworthy and poorly known species found near the Rio Grande do
Sul border is aimed at improving the knowledge of gasteroid fungi in Santa
| Catarina State.
298 ... Cortez & al.
Materials and methods
Specimens were collected in the State of Santa Catarina, southern Brazil, near
Rio Grande do Sul border, and kept at the SMDB herbarium. Morphological
studies were performed according to Miller & Miller (1988). The color
terminology used is that of Kornerup & Wanscher (1978). Line drawings of the
microstructures were made with the aid of a drawing tube. Scanning electron
microscopy (SEM) studies were performed with a Phillips XL30; basidiospores
were mounted on aluminum stubs and coated with 10-15 nm of gold using a
standard sputter.
Taxonomy
Geastrum ovalisporum Calonge & Mor.-Arr.,
Bol. Soc. Micol. Madrid 25: 273, 2000. FIG. 1-4
Basidiomata 30-36 mm high, 22-26 mm diam, epigeous at maturity (eggs
not seen). Exoperidium fornicate (KW 5F5), non-hygroscopic, splitting in 7
unequal rays; mycelial layer encrusted with debris; fleshy layer brown (KW
5F4), sloughing away to reveal a fibrous layer. Endoperidial body 12-14 mm
diam, subglobose, dark brown (KW 7F4), pedicellate (2-3 mm high) and with
a distinct hemispheric apophysis. Peristome fimbriate to slightly striate and
conical, well delimited by a circular and slightly depressed area in the centre
of the endoperidium, dark brown (KW 6F4). Gleba pulverulent when mature,
dark brown (KW 7F4).
Fic. 1-3. Geastrum ovalisporum.
1. Basidioma. 2. Basidiospores. 3. Capillitium.
Gasteroid fungi from Santa Catarina (Brazil) ... 299
Basidiospores 3-3.5 x 2-2.5 wm diam (incl. ornamentation), ovoid to
subglobose, pale brown under KOH, with a small verrucose ornamentation,
short tubular pedicel present, under SEM the ornamentation is strongly
verrucose. Capillitium formed by 2-5 wm diam hyphae, with yellowish,
thickened walls, lumen much reduced or absent.
EXAMINED SPECIMEN: BRAZIL. Santa Catarina State, municipality of Mondai, 12.X.1995,
leg. A.A. Spielmann (SMDB 11.522); on soil and litter in subtropical forest.
Discussion: Calonge et al. (2000) described Geastrum ovalisporum with
the following diagnostic characters: non-hygroscopic exoperidium, stipitate
endoperidium, fimbriate peristome, and ovoid basidiospores, which are
unique in the genus. The collected specimens were found growing on litter
and subtropical rainforest soil, in contrast to the Bolivian holotype, which was
collected on sandy soil in tropical forest. Leite et al. (2007) recently reported
G. ovalisporum for the first time from Brazil (in Rio Grande do Norte State) on
soil with woody debris. ‘This is the first report of G. ovalisporum from southern
Brazil.
Fic. 4-5. Basidiospores under SEM.
4. Geastrum ovalisporum. 5. Tulostoma rickii.
_ Tulostoma rickii Lloyd, The Tylostomeae: 20, 1906. Fic. 5-8
| = T: bulbillosum Bres., Broteria 5: 28, 1906.
| Basidiomata 56-81 mm high, epigeous. Endoperidial body 18-23 mm diam,
| subglobose. Exoperidium membranous, light orange (KW 5A5) to yellowish
brown (KW 5D8), encrusted with soil particles and thus with a verrucose
| surface. Endoperidium membranous, light orange (KW 5A4), adhered to the
exoperidium. Peristome fibrillose, almost indistinct. Socket conspicuous to
| inconspicuous, with a membranous consistency, scaly. Gleba pulverulent when
mature, brownish yellow (KW 5C7) to golden brown (KW 5D7). Stipe 42-71
x 5-7 mm, slightly sinuous, with longitudinal striae, scaly surface, light orange
(KW 5A5) to yellowish brown (KW 5D6), base volviform with abundant soil
particles adhered, hollow with white flesh.
300 ... Cortez & al.
Fig. 6. Basidiomes of Tulostoma rickii.
Basidiospores 4.2-5 wm diam (excl. ornamentation) or 6.7-8.4 “m (incl.
ornamentation), globose, distinctly clathrate and appearing with ‘wings or false
‘spines’ in optical section. Capillitium hyphae 4.2-8.4 vm diam with thickened
walls, hyaline to stramineous, septate.
EXAMINED SPECIMEN: BRAZIL. Santa Catarina State, municipality of Riqueza,
27.X11.2006, leg. A.A. Spielmann & M.A. Sulzbacher 86 (SMDB 10.999); growing on
humic soil, among roots, in subtropical rain forest.
Gasteroid fungi from Santa Catarina (Brazil) ... 301
Fic. 7-8. Tulostoma rickii.
7. Basidiospores. 8. Capillitium.
Discussion: The following Tulostoma species have been reported from Santa
Catarina State: T’ cyclophorum Lloyd and T! exasperatum Mont. (Wright
1987). Tulostoma rickii is diagnosed by the presence of a bulbous stipe base
that resembles a pseudovolva, and reticulate basidiospores (Wright 1987).
The basidiospores are similar to those of T: exasperatum, which is, however,
lignicolous in contrast to T. rickii, which occurs on rainforest soils. Wright
(1987) also considered T. rickii a rare Tulostoma species reported only from
Rio Grande do Sul in Brazil and from Argentina. This is the first report from
Santa Catarina.
: Acknowledgements
The authors thank Adriano Spielmann for help in collection. Special thanks to Dr.
Francisco de Diego Calonge (Real Jardin Botanico, Madrid, Spain) and Dr. Maria Alice
Neves (Universidade Estadual de Feira de Santana, Brazil) for critical review of the
“manuscript. CNPq (Brazil) is acknowledged for financial support.
Literature cited
|
Calonge FD, Moreno-Arroyo B, Gomez J. 2000. Aportacion al conocimiento de los gasteromycetes,
Basidiomycotina, de Bolivia (América del Sur). Geastrum ovalisporum sp. nov. Bol. Soc. Micol.
| Madrid 25: 271-276.
| Fidalgo O. 1965. Introduc4o a historia da micologia brasileira. Rickia 3: 1-44.
Giachini AJ, Oliveira VL, Castellano MA, Trappe JM. 2000. Ectomycorrhizal fungi in exotic
| Eucalyptus and Pinus plantations in southern Brazil. Mycologia 92: 1166-1177.
| Kornerup A, Wanscher JH. 1978. Methuen handbook of colour. 3™ ed. London (UK): Eyre
Methuen.
302 ... Cortez & al.
Leite AG, Calonge FD, Baseia IG. 2007. Additional studies on Geastrum from northeastern Brazil.
Mycotaxon 101: 103-111.
Miller Jr. OK, Miller HH. 1988. Gasteromycetes: Morphological and Development Features With
Keys to Orders, Families, and Genera. Eureka (USA): Mad River. 157 p.
Moller A. 1895. Brasilische Pilzblumen. Jena (Germany): Gustav Fischer. 152 p. + 8 pl.
Sobestiansky G. 2005. Contribution to a macromycete survey of the States of Rio Grande do Sul
and Santa Catarina in Brazil. Braz. Arch. Biol. Technol. 48: 437-457.
Viégas AP. 1945. Alguns fungos do Brasil X. Gastromicetos. Bragantia 5: 583-595.
Wright JE. 1987. The genus Tulostoma (Gasteromycetes) — a world monograph. (Bibliotheca
Mycologica 113). Berlin/Stuttgart (Germany): J. Cramer. 338 p.
MYCOTAXON
Volume 106, pp. 303-309 October-December 2008
Validation and typification of the name Claviceps zizaniae
S.A. REDHEAD’, M.E. CORLETT & M.N.L. LEFEBVRE
redheads@agr.gc.ca
‘National Mycological Herbarium
Eastern Cereal & Oilseed Research Centre, C.E.F., Agriculture & Agri-Food
Canada, Ottawa, Ontario, Canada, KIA 0C6
Abstract — The binomial, Claviceps zizaniae, has not previously been validly published.
The basionym, Spermoedia zizaniae, was improperly neotypified and is here lectotypified
and that type is epitypified. The history of the discovery and description of this ergot
fungus on wild rice, Zizania aquatica, and the roles of the main contributors, Faith
Fyles, Charles M. Wright, Ibra Conners and Maria Pantidou, are outlined.
Key words — Cornell, DAOM, nomenclature, sclerotia, teleomorph
Introduction
Claviceps zizaniae is the commonly used name for the fungus forming ergot
on the wild rice species, Zizania aquatica L. and Z. palustris L. (Aiken et al.
1988, Conners 1967, Dore 1969, Farr et al. 1989, Ginns 1986, Kernkamp et al.
1976). Phylogenetically, C. zizaniae continues to be an accepted, well defined
taxon (Alderman 2003, Alderman et al. 2004, Pazoutova et al. 2004, 2008).
Data-basing of archived specimens in the Canadian National Mycological
Herbarium (DAOM, see Holmgren & Holmgren 1998) revealed the existence of
both a purported ‘neotype and ‘original material’ for the basionym, Spermoedia
zizaniae Fyles (Fyles 1920b). Further investigation led to the realization that
the basionym should not have been neotypified (Pantidou 1959) and that the
combination in ‘Claviceps zizaniae (Fyles) Pantidou’ (Pantidou 1959) has been
invalid until now.
History
Ergot on wild rice in North America has long been known to aboriginal
peoples, such as the Ojibwa, who apparently referred to it in words meaning
“frozen rice” (Jenks 1901) or “frozen kernels” (Denniston 1900). The earliest
record in technical literature was by Trelease (1886), who differentiated the
'© Her Majesty the Queen in Right of Canada, as represented by the Minister of Agriculture and
Agri-Food Canada
304 ... Redhead, Corlett & Lefebvre
ungerminated, pinkish brown, stout, ovoid sclerotia of Claviceps sp. on Zizania
aquatica in Wisconsin from common ergot (Claviceps purpurea (Fr.) Tul.) on
other grains. The next report in the literature by Farlow & Seymour (1888) is
traceable to Trelease’s report via the publication by Farlow & Trelease (1887).
Unaware of Trelease’s 1886 report, Fyles (1915) cited Denniston’s 1900 report of
Claviceps purpurea ergot on Zizania aquatica as the earliest record. Fyles (1915)
was the first to overwinter and induce germination of the sclerotia in order
to view the characteristically stalked stromatal heads with fertile perithecia
and asci characteristic of a Claviceps. Additionally, she performed inoculation
studies demonstrating that the Claviceps on Zizania only infected Zizania and
did not infect normal hosts for C. purpurea (that she called Spermoedia clavus
(DC.) Fr.), i.e. Agropyron tenerum Vasey (= Elymus trachycaulus (Link) Gould ex
Shinners), Alopecurus pratensis L., Arrhenatherum elatius (L.) P. Beauv. ex J. Presl
& C. Presl, Dactylis glomerata L., Poa pratensis L. and Elymus dasystachys Trin.
(= Leymus secalinus (Georgi) Tzvelev). Notably, Fyles (1915) also mentioned
that the sclerotia from wild rice floated, which is biologically significant for a
swamp inhabiting fungus (Alderman et al. 2004, Brown 1947). At the end of her
publication, Fyles (1915) promised to provide a formal description and name
for the fungal species elsewhere. As noted by Pantidou (1959), Fyles never did
provide a more formalized description, however in her subsequent bulletin on
wild rice, Fyles (1920b) repeated some of the information and photographs,
and on page 17 named the fungus Spermoedia zizaniae with a further citation
of her earlier publication in a footnote. Publication of the species in Spermoedia
Fr. rather than in Claviceps Tul. was, as commented upon by Pantidou (1959),
undoubtedly influenced by the classification published by Seaver (1911), who
treated Claviceps as a synonym of Spermoedia. In her bulletin on poisonous
plants, Fyles (1920a) adopted the name Spermoedia clavus with Claviceps
purpurea in synonymy, when she described ergot of rye. Also revealing her
mindset was the fact that in 1915 she had stated that Claviceps microcephala
(Wallr.) Tul. was “now” called Spermoedia microcephala (Wallr.) Seaver.
Pantidou (1959) correctly noted that Art. 59, then, and even today (McNeill et
al. 2006: 59.2 & Ex 5), leads to the interpretation of Fyles’ name, “Spermoedia
zizaniae’, as a teleomorphic name because she included a description of the
teleomorph complete with descriptions of asci, ascospores and perithecia. It
was Pantidou'’s work, done in collaboration with Dore who was working on
wild rice (Dore 1969), that precipitated modern acceptance of the fungus as a
distinct species. Unfortunately, the combination, Claviceps zizaniae proposed.
by Pantidou (1959) was invalidly published, in contravention of Art. 33.4.)
Although Pantidou cited Spermoedia zizaniae Fyles (1920b) as the basionym,|
she failed to provide a full and direct reference to the place of publication)
(Fyles 1920b: 17). Fyles’ (1920b) publication, where the name was validated,)
Validation of Claviceps zizaniae ... 305
was a discussion on wild rice and therefore cannot be interpreted as publication
coextensive with the protologue for the fungal name (Art. 33, Note 1). Technical
validation is provided below.
Fic. 1. Slide prepared by C. Wright from original Fyles material of
Spermoidea zizaniae DAOM 12023.
Pantidou (1959) also designated a ‘neotype’ for the name (DAOM 56855 [=
Dore # 17042]), which was further illustrated in the publication by Dore (1969:
62, as Dore # 17042, in situ). Although this proposed typification has provided
stability to the usage of the name C. zizaniae, designation of it was also an
error, because it consisted of two elements, the original sterile sclerotia and
perithecia induced 2 years later. A neotype may only be designated when no
original material exists (Art. 9.6). Illustrations, such as those provided by Fyles
(1915, 1920b) can be designated as types (Art. 9.2) and can constitute original
material as defined by Art. 9, Note 2. Hence, the designated ‘neotype’ is to be
superseded (Art. 9.17). Additionally, there is other original material to select
from as is explained below.
Faith Fyles was an assistant botanist under Hans Gtissow, the Dominion
Botanist at the Department of Agriculture (Lloyd 1961) in Ottawa. Some of
her materials remained with the department in Ottawa. For some years, the
importance of her work demonstrating host specificity was underestimated by
others who went on to identify ergot of Zizania as Claviceps purpurea (Conners
1931, 1938, Sprague 1950, Wehmeyer 1950).
In 1940, the wild rice ergot was discovered on Zizania in Maine, USA, by the
botanist, Steinmetz (1940), leading to a student's Master’s thesis (Wright 1942)
anda published abstract fora meeting of the American Phytopathological Society,
which, however, was cancelled because of World War II (Steinmetz & Wright
1943). The teleomorph of Claviceps zizaniae was once again germinated from
sclerotia and a dipterous vector, Oscinella neocoxendix Sabrosky [now Apallates
neocoxendix (Sabrosky)], was identified (Steinmetz & Wright 1943). The host
specificity was confirmed, i.e. the ergot fungus from Zizania would only infect
306 ... Redhead, Corlett & Lefebvre
Zizania and not barley, oats, wheat, or millet, while Claviceps from Festuca,
Calamagrostis, Agropyron, and Dactylis would not infect Zizania. Evidently the
latter publication led to correspondence between Ibra Conners, in Ottawa, and
C. M. Wright, then a student of Prof. Patrick Fitzgerald at Cornell University.
In a letter filed in DAOM with specimens, dated July 30, 1944, Wright thanked
Conners for sending Fyles’ material, noting that he was returning it undivided
at the suggestion of Fitzpatrick, and that he was including a slide of a macerated
perithecial head plus his germinated sclerotium from the material in Maine.
On file in DAOM are the specimen of germinated sclerotium sent by Wright
(DAOM 239767) and the slide of the macerated perithecial head he prepared
from Fyles’ original specimen (DAOM 12023), specifically noting on the slide
label “Ottawa Herbarium 12023”. The actual germinated sclerotium/sclerotia
with perithecial heads that Fyles labeled Spermoedia zizaniae and which were
sent to Wright cannot be located in DAOM. However, the DAOM accession
book records for 12023 “on Zizania aquatica L. Ottawa June 30, 1914 FE Fyles
EF” Notably, June 1914 was the last month that Fyles (1915) recorded for
maturation of the perithecia on germinated sclerotia collected in Ottawa and
overwintered in Ottawa. Hence, the squash mount of DAOM 12023 (Fig. 1) is
considered to be original material. However, given its poor state of preservation,
we prefer to designate the illustration (unnumbered plate on p. 18, Fyles 1920b)
as lectotype, rather than the slide, and further to choose the portion of the |
‘neotype’ selected by Pantidou (1959) that bears perithecia [designated as
DAOM 56855(b)] as the epitype to the lectotype. We purposely do not choose
pl. XI from Fyles (1915) because it is a collage of photographs, possibly from
different dates, whereas the 1920 photograph constitutes a single photographic
plate. We differentiate the original collection of sterile sclerotia collected by
Dore as DAOM 56855(a) — see below.
Charles Wright had indicated in his letter to Ibra Conners that he intended to
publish the combination in Claviceps in Mycologia but for unknown reasons,
no such combination was made in Mycologia (or elsewhere) nor did any
publication by Wright concerning Claviceps ever appear in Mycologia. Instead
he went on to study viral diseases in tomatoes and cherries (Wright 1947, |
Blodgett & Wright 1948). Of historical significance is a specimen by Wright
(CUP 31718) and photographs (CUP 31717, 31719-31721) deposited at Cornell
University bearing labels with the name “Claviceps zizaniae (Fyles) Wright”
(K. Hodge, curator CUP, pers. comm. 2008) and the slide of Fyles’ specimen
he returned to DAOM bore the binomial “Claviceps zizaniae”. However, not
until Pantidou (1959) published her paper on Claviceps zizaniae did the species
became generally accepted as distinct from C. purpurea by Conners (1967) and |
later authors and host indices compilers. Then the name Claviceps zizaniae —
came into usage.
Validation of Claviceps zizaniae ... 307
In order to preserve this oft-cited link to the literature, we propose the following
combination.
Nomenclature
Claviceps zizaniae (Fyles) Pantidou ex Redhead, M.E. Corlett & M.N.L. Lefebvre,
comb. nov. FIG. 1
MycoBANnkK MB512148
BASIONYM: Spermoedia zizaniae Fyles, Wild rice. Dominion Experimental Farm,
Canad. Dept. Agric., (2" ser.) Bull. 42: 17. 1920.
“Claviceps zizaniae (Fyles) Wright’, nom. herb. (CUP), nom. inval. (Art. 29.1).
“Claviceps zizaniae (Fyles) Pantidou’, Can. J. Bot. 37: 1234 (1959), nom. inval.
(Art. 33.4).
LectroryPus for Spermoedia zizaniae (here designated): Dominion of Canada. Dominion
Experimental Farm, Dept. Agric., Bull. 42 (2° ser.): unnumbered photograph (p. 18)
(Fyles 1920b) {teleomorph}.
Photographed material, although published in 1920, is presumed to represent
overwintered sclerotia originally harvested with seed of Zizania aquatica, Sept. 29, 1913,
Treadwell [45° 35° 49” N - 75° 00’ 38” W], Ontario, Canada, and variously treated in
Ottawa to induce germination as outlined by Fyles (1915). Fyles (1920b) indicated that
work on wild rice, itself, was halted after her 1913-14 research until 1917 after which year
wild rice access was indirectly curtailed because gatherers had gone to the 1“ world war
front [1914-18]. The only other collection associated with Fyles in DAOM was DAOM
35777, on wild rice from Shediac, New Brunswick, Canada, Oct. 14, 1914 [“14/10/14"]
based upon sclerotia only, and originally labeled Claviceps purpurea in pencil, written
over in Indian ink, with the species epithet crossed out and replaced by “zizaniae (Fyles)
Pantidou’, obviously relabeled concurrently with or after Pantidou’s publication.
Epitypus (here designated): DAOM 56855(b), harvested 20 Feb. 1959, perithecia on
stroma germinated after soaking sclerotia in sterile water in June 1958, refrigerated
until November, and transferred to sterile sand in greenhouse, using sclerotia originally
collected on Zizania aquatica var. interior, in swamp woods, about 2 miles NW of
Sheffield, New Brunswick, Canada, 9 Sept. 1957, W.G. Dore 17042 [sclerotia filed as
DAOM 56855(a)]. See Pantidou (1959) for additional details.
Acknowledgements
We thank Dr. Kathie Hodge for researching literature and specimens by Charles Milton
Wright at Cornell University (CUP) and reviewing the manuscript We also thank Dr.
Amy Rossman (USDA, BPI) for presubmission review.
Literature cited
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Toronto.
Alderman SC. 2003. Diversity and speciation in Claviceps. p. 195-246, in White JE, Bacon CW,
Hywel-Jone NL, Spatafora JW. [eds.] Clavicipitalean Fungi: evolutionary biology, chemistry,
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Alderman SC, Halse RR, White JE. 2004. A reevaluation of the host range and geographical
distribution of Claviceps species in the United States. Plant Dis. 88: 63-81.
308 ... Redhead, Corlett & Lefebvre
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Jenks AE. 1901. The wild rice gatherers of the Upper Lakes. U.S. Dept. Int., Bur. Amer. Ethnol., 9"
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126-135.
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grohii: Their Alkaloids and Phylogenetic Placement. J. Nat. Prod. - por 10.1021/np8001173
Seaver FJ. 1911. The Hyporeales of North America: IV. Tribe IV. Cordycipiteae. Mycologia 3:
207-230. ;
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Univ.
MYCOTAXON
Volume 106, pp. 311-360 October-December 2008
Revision of Scutel/lospora and description of
five new genera and three new families in the
arbuscular mycorrhiza-forming Glomeromycetes
FRITZ OEHL', FRANCISCO ADRIANO DE SOUZA?” & EWALD SIEVERDING?
'Zurich-Basel Plant Science Center, Institute of Botany, University of Basel
Hebelstrasse 1, CH-4056 Basel, Switzerland
?Embrapa Agrobiologia, Seropédica
Rodovia BR465 km 7, CEP 23890-000, Rio de Janeiro, Brasil
**sieverdingE@aol.com
3Institute for Plant Production and Agroecology in the Tropics and Subtropics
University of Hohenheim, Garbenstrasse 13, D-70599 Stuttgart, Germany
Abstract — Arbuscular mycorrhizal fungi forming spores on bulbous sporogenous
cells were earlier organized in two genera of the Gigasporaceae: Gigaspora and
Scutellospora. Molecular analyses indicated that Scutellospora is polyphyletic. The
Gigasporaceae are revised on the basis of morphological spore characters and 18S and
25S rRNA gene sequences, and the 36 Scutellospora species are reorganised in three
new families including five new genera: Scutellosporaceae (Scutellospora), Racocetraceae
(Racocetra, Cetraspora) and Dentiscutataceae (Dentiscutata, Fuscutata, Quatunica).
The family Gigasporaceae remains with a single genus Gigaspora. The molecular data
indicate that the genus Gigaspora derived from within the former genus Scuteilospora.
The family Scutellosporaceae forms the most ancestral clade, while Gigasporaceae are
phylogenetically not much distant from Racocetraceae and Dentiscutataceae. Keys are
presented for the identification of all species of the reorganized former Gigasporaceae.
Key words — Glomeromycota, Diversisporales, germination shield, ontogeny,
phylogeny
Introduction
| Arbuscular mycorrhizal fungi (AMF), which form spores terminally on
| bulbous sporogenous cells, were described in the family Gigasporaceae (Morton
& Benny 1990) of the order Diversisporales, Glomeromycetes (Cavalier-Smith
| 1998). In this family, with about 40 species, spores are relatively big, generally
| >200 um and up to 700-1050 um in diameter. Besides having this typical spore
formation, species of the Gigasporaceae also differ morphologically from other
312 ...Oehl, de Souza & Sieverding
families of the Glomeromycetes by not forming intraradical vesicles but instead
so-called auxiliary cells in the extraradical mycelium (Gerdemann & Trappe
1974, Morton & Benny 1990). Two genera have been taxonomically recognized
based on spore wall characteristics and germination structures: Gigaspora
and Scutellospora (Walker & Sanders 1986, Morton & Benny 1990). Spores of
Gigaspora spp. differentate a single spore wall and multiple germ warts on the
inner wall layer (= single wall group sensu Walker 1983), and Scutellospora spp.
form two to four spore walls and a discrete germination compartment (the
so-called germination shield) on the innermost wall. The spore wall structure,
spore ontogeny and development of the mycorrhizal mycelia in Scutellospora
were precisely described by Franke & Morton (1994), Morton (1995), de Souza
& Declerck (2003), Declerck et al. (2004), de la Providencia et al. (2005) and de
Souza et al. (2005).
The last publication unequivocally showed through molecular-based
phylogenetic analyses that Scutellospora is polyphyletic, a finding that was
also evident in other recent articles (e.g., Walker et al. 2004, Ahulu et al. 2006,
Redecker et al. 2007). However, de Souza and co-workers found a disagreement
between morphological based and molecular cladistic analyses that they could
not resolve.
The objective of this study was to look at common morphological features of the
spores of all species in the genus Scutellospora and to search for morphological
conformity and differences, and to find common morphological features of —
spores for the species that were congruent with the rRNA-based molecular —
phylogenetic reconstruction. We concentrated our investigation on spore
morphology because spores are the basis for species diagnosis in AMF. We had
access to several collections of spore specimen of Scutellospora spp. Specimens
with mycelium and in particular auxiliary cells in the root external mycelium
were seldom available from species described of the Gigasporaceae, so that we —
did not consider these fungal parts in our investigations. According to Walker
& Sanders (1986) the morphology of the auxiliary cells does not show much —
variability in Scutellospora anyway. |
We analyzed type or isotype specimens of almost all described Scutellospora spp.
and present germination shields of 31 species in colored photographs. We found |
that there are common morphological spore features in Scutellospora that are ©
congruent with the molecular phylogenetic groups based on rRNA genes. The |
corresponding morphological and molecular groups are hereafter presented |
first. According to these groups, we then emend the family Gigasporaceae, ©
reorganize the currently known species of Scutellospora taxonomically into ©
three new families and six genera of the Diversisporales, and transfer 25 species |
of the Scutellospora genus into five new genera. :
New taxa in the Glomeromycetes ... 313
Material and methods
Specimen analyses
The Scutellospora specimens analyzed, their provenance and locations are listed in
TABLE 1. In particular ex type, isotype and paratype material were examined, and we
had access to several herbaria and private collections of arbuscular mycorrhizal fungi
(e.g., OSC, FH, URM (Recife, Brazil), Embrapa Agrobiologia (Seropédica, Brazil), DCS-
UFLA (Lavras, Brazil), International Culture Collection of Arbuscular and Vesicular-
Arbuscular Mycorrhizal Fungi (INVAM), and personal collections held by Sieverding,
Oehl, de Souza, Trappe, Spain, Blaszkowski, Cuenca, and C. Castillo). Additionally, all
original species descriptions and published species emendations were also considered.
During a visit at INVAM, F. Oehl analyzed stored vouchers of the following species:
S. arenicola, S. biornata, S. calospora, S. cerradensis, S. coralloidea, S. dipurpurescens,
S. erythropus, S. fulgida, S. gregaria, S. heterogama, S. pellucida, S. persica, S. reticulata,
S. rubra, S. scutata, S. verrucosa, S. weresubiae. Spores of the following species were
isolated from living culture accessions at INVAM and permanently fixed on microscope
slides before analyzing: S. calospora, S. coralloidea, S. dipurpurescens, S. erythropus,
S. fulgida, S. heterogama, S. persica, S. verrucosa.
Older specimens (mounted on microscopic slides before 1990) had generally been
mounted either in lactophenol, while other specimens were fixed with polyvinyl alcohol-
lactic acid-glycerol (PVLG) or in a mixture of PVLG + Melzer’s reagent, which has
become the major fixing media since 1990 (e.g. Brundrett et al. 1994). We fixed newly
mounted spores from the collections using those two recent mountant media. When
available, spores freshly isolated from soils were also mounted in PVLG or in PVLG +
Melzer’s reagent and sometimes also in a mixture of lactic acid to water at 1:1, in Melzer’s
reagent, and in water. Spore walls, germination structures and all other mycorrhizal
‘structures were analysed using compound microscopes at 100-630x magnification.
Most photographs presented in this paper were taken with a digital camera (Olympus
‘model DP70-CU) on a Zeiss Axioplan compound microscope. A few photographs were
taken by F. Oehl at INVAM or at OSC. The legends and scales on the photographs of the
figures were inserted with Adobe Photoshop CS2 9.0.
Spore wall terminology
The terminology of the spore walls in our paper differs from the nomenclature
for walls followed by Morton (1995) and Silva et al. (2006b) in Scutellospora (see
also INVAM’s homepage: www.invam.caf.wvu.edu). INVAM and Silva et al. (2006b)
| differentiate two to four walls: the spore wall (SW) and one to three germinal walls
|(Gw1, Gw2 and Gw3), each with different layers. This nomenclature is misleading
because in the Gigasporaceae spore germination always starts from the innermost layer
| of the innermost wall, from which we conclude that spores have only one germinal wall,
|and not several. Thus, we adopt the wall terminology we used for other AMF genera
\(e.g. Oehl & Sieverding 2004, Oehl et al. 2006, Spain et al. 2006, Palenzuela et al. 2008)
|\for Scutellospora species: INVAM’s ‘spore wall’ (sw) is termed here the ‘outer spore wall’
(ow) while INVAM’s cw, (where n = 1-3) we call ‘middle wall(s)’ (mw,, where n=1-2)
and ‘inner wall’ (tw). Our Iw always is the germinal wall in all Scutellospora spores, as
a germination shield is formed on the surface of this wall or between a thin outer layer
jand subsequent layers of the 1w.
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(edojorqo8y edeiquiyg pue yTAN -SOd)
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(ZUIpIOAaIS) eIQUIO[OD WOT] suaumtoeds
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(Y20) WVANI X2 ‘AVANT 38 suouttseds
({yaO) puepeziMs pure AueUtIay
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(BUIPIOASIS) BIQUIO[OD WOT} suauriseds
(eznos ap) [Izerg Worf pure WVANI 3 suatads
(T4290)
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(TY9O) puelaziIMs
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(1420)
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Hal S0S adAiost ‘77,974 OSO edAjofOH
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318 ...Oehl, de Souza & Sieverding
Phylogenetic analyses
The phylogenetic relationship of species in the Gigasporaceae was reconstructed by
independent analyses of 18S and 25S rRNA genes. Nearly complete sequences of 15
Scutellospora spp., seven Gigaspora spp. and one Pacispora were used to reconstruct the
18S rDNA-based phylogeny. Pacisporaceae sequences were chosen as outgroup because
it is the closest family to Gigasporaceae (Walker et al. 2004). For the 25S rRNA genes
partial sequences of 8 Scutellospora and 3 Gigaspora species were used. In this case, the
Acaulosporaceae was selected as outgroup, as they were the closest group with sequences
available (Silva et al. 2006a). The majority of the sequences were retrieved from public
databases, with exception of some Gigaspora 18S rDNA sequences that were updated
from previous work (de Souza et al. 2004). The sequences were aligned automatically
using Clustal X (Thompson et al. 1997). The alignments were further improved by
visual inspection and edited using SeaView (Galtier et al. 1996). The phylogeny was
reconstructed by Distance, Maximum Parsimony, and Maximum Likelihood criteria
using PAUP* version 4.0 Beta 10 (Swofford 2003). For distance (minimum evolution)
and maximum likelihood criteria the trees were obtained using heuristic search with
likelihood settings from best-fit model selected by Akaike Information Criterion in
ModelTest 3.7 (Pousada & Crandall, 1998) and for Maximum Parsimony, the trees were
obtained using two ways (1) considering gaps as fifth state or (2) as missing characters:
The bootstrap analyses were carried out using 1000 replicates, plus 10 additional
replicates per run, by heuristic search with sequences added randomly and branch
swapping “tree-bisection and reconnection” as best tree search method. The Gigaspora
sequences were updated from previous entries (see PLATE 2 for accession numbers) and
the alignments generated in this study were deposited in EMBL-EBI nucleotide sequence
database (http://www.ebi.ac.uk) under the alignment numbers ALIGN_001298 and
ALIGN_001257.
Results
General morphological spore analyses
In order to identify corresponding morphological differences between the
genetically different groups (groups A, B and C according to de Souza et al.
2005), we first listed all major spore characteristics of the Scutellospora spp.
(TABLE 2). Spore size, spore and wall colors, wall ornamentations, numbers of
walls, numbers of wall layers, staining features of wall layers and morphology of —
the germination shield alone were not suitable characters for grouping spores
with the molecular clades. Only the number of spore walls combined with the
germination shield morphology could be related to the genetically different
groups. Additionally, we noticed that the positive staining reaction of the inner
wall in Melzer’s is somehow linked with the number of spore walls. According to
these three major morphological characters, we sub-divided de Souza’s groups
B and C into sub-groups (B1, B2 and B3, and Cl and C2, respectively). Since
germination shield morphology is so important for diagnosing the different
groups, we describe their formation and their differences in the following.
New taxa in the Glomeromycetes ... 319
| PLATE 1— FIGS. 1-9: CHARACTERISTICS OF TYPICAL GERMINATION SHIELDS. All shields have
i) a germination hole (gh) bridging in/on the innermost wall between the spore cytoplasm
and the germination shield, ii) a variable number of germ tube initiations (gti) from where
| germination tubes may start growing, and iii) folds on the shield separating into different lobes and
compartments, respectively. FIGs. 1-4. Bi-lobed shields: 1. Violin-shaped. 2. Oval to ovoid shaped.
3. Bi-lobed, with a few folds on the shield wall. 4. Mono-lobed, coiled. Frc. 5. Open organized,
|wavy shield with multiple small compartments; wavy compartments separated by folds. Each
|compartment generally has one central gti from where the germ tubes emerge during germination.
FIGS. 6-9. Compartmented shields with several to many (8-30) small compartments, separated
by folds; each compartment having one gti in its center. Shield shapes circular, cardioid, reniform
or ellipsoid. Fics. 8-9. Highly complex structured germination shields; shield peripheries are
conspicuously denticulate.
320 ...Oehl, de Souza & Sieverding
Morphology of germination shields
Common to all Scutellospora spore germination shields (also called “germ
shield”) is their formation on the inner wall facing the next outermost wall (mw
Or OW, see TABLE 2). Very seldom does the literature report two germ shields on
the same spore. All germ shields initiate from a germination hole (gh) through
the innermost Iw layer or — more generally —- through the innermost spore wall
IW (PLATE 1). Walker & Sanders (1986) described the germ shield formation
process as an extrusion, with the plasma membrane extending through the hole
to form lobes on the rw surface or between a thin Iw outer layer and subsequent
adherent Iw layer. The germ shield size ranges from ca. 35 x 70 um to about 140
x 160 ttm and up to 180 x 280 um. The germ shield may be divided into one,
two, several, or multiple (up to 30) small compartments (sensu Gerdemann &
Trappe 1974) that are described below. Each compartment can contain a small,
more or less circular (2-5 um to 12-20 um in diameter) germ tube initiation
(gti, Walker & Sanders 1986, PLATE 1), from which a germination tube starts
growing through the overlaying wall layers. Generally, however, only one or
two (of 30 gti) actually form a germ tube during germination. Once grown
through the walls, the germ tube may begin to branch on or near the spore
surface although it often grows straight into the soil. Germ shields are rather
more frequent (or at least more easily seen) on older rather than young spores,
probably because the shield forms at the end of spore wall differentiation
(Franke & Morton 1994, Morton 1995, de Souza et al. 2005). On some species
germ shields are readily visible on young mature spores (e.g. S. dipurpurescens) |
whereas they are almost never visible on others (e.g. S. pellucida). It should |
also be noted that, in a very young or early stage of germ shield formation, —
gti are not visible, suggesting that they may form later in the shield formation :
process. |
In principle, we differentiate the germ shields morphologically by color, |
spherical shape (including dentate structures), and number and shape of lobes |
or compartments, each with its own gti. In different species, germ shield size
is generally related to spore size and is thus so variable that it is not a good
diagnostic character for identifying species groups.
Germ shield color
Three of the genetically different groups, A, Cl and C2, form transparent |
hyaline to very light yellowish germ shields, e.g. S. calospora, S. coralloidea \
and S. pellucida (PLATES 4-6), whereas the genetic groups B1, B2 and B3 form
distinctive yellow brown to dark brown pigmented germination shields, e.g.
S. heterogama, S. nigra and S. erythropus (PLATES 7-9). The pigmentation of the
germ shield is not correlated to the color of the spore walls; for example, the’
shield is brown-yellow to brown in white-spored S. cerradensis and S. scutata)
New taxa in the Glomeromycetes ... 321
but hyaline to light yellow in S. coralloidea whose spores are dark colored. Some
hyaline germ shields, when mounted on slides or stored in liquids of different
media for several years, may somewhat darken and appear brownish: e.g., a
germ shield of S. coralloidea isolated from the type at OSC that had been stored
in lactophenol for 35 years had a partly darkened germ shield wall (Frc. 20).
Likewise, the inner spore wall including the germ shield of S. tricalypta was
darkened (FIG. 14), 25 years after being fixed on a slide in Ferrant mountant by
Ferrer & Herrera (1981).
Germ shield shape
In planar view, germ shields are violin-shaped, oval to ovoid, coiled, circular,
cardioid to reniform, or ellipsoid (PLATE 1). In principle three morphologically
different germ shield types are observed (see TABLE 2, PLATES 1 & 4-9). Species
of the genetic groups A have simple shields that are bi-lobed (Fics. 1-3) — with
the inner borders between the folds often forming a lyre, as in S. calospora — or
(rarely) mono-lobed (FIG. 4) and coiled, as in S. projecturata. Such shields have,
near the end of each lobe, one gti. In some specimens, simple bi-lobed shields
appeared to have three or four lobes but these ‘additional’ lobes were only
prolongations of the two initial lobes, or infolds of the same lobes that became
apparent when the shield could not be observed in completely planar view (e.g.,
FIGS. 3, 18 for S. nodosa). Group B1 germ shields are also rather simple, bi-
lobed and lyre-like (e.g. S. heterogama and S. rubra, Fics. 45-47); in group B1
as well, further infolds of the two initial lobes may sometimes cause the shield
to appear to have three or four lobes (e.g. S. savannicola in F1G. 48) even though
group Bl germ shields have only (1-)2 gti. Species assigned to genetic groups
Cl and C2 have a distinctly different germ shield morphology and a clear lyre
shape is not visible. Multiple wavy lobes form an open network of many small
compartments with up to 12 compartments with respective gti visible (FIG. 5 +
PLATES 5-6). Finally, genetic groups B2 and B3 have highly complex-structured,
compact shields with from 8-10 to 30 compartments, each with each one gti
(FIGS. 6-9). The shield periphery of these species generally looks conspicuously
dentate (e.g. FIGS. 8-9 + PLATES 7, 9).
Spore wall numbers
| The outer wall (ow) in all Scutellospora spp. has three to four layers and
| the inner wall (1w) generally is 2-to-3-layered. Differences between the genetic
| groups are found in the number (0-2) of middle walls (TABLE 2). Genetic group
|Cl1 species lack any Mw. Group A, B1, B2 and C2 species have one single- or
bi-layered middle wall (mw; called GW1 by Morton 1995). Only, S. erythropus
|(group B3) forms two middle walls (mw1 and Mw2).
Staining reaction of spore walls
Scutellospora spp. with one or two middle walls (i.e. generally all species in
morphological groups A, B1, B2, B3 and C2) stain purple to deep purple in
Melzer’s reagent on the hyaline inner wall. In these groups, Melzer’s staining
reactions are less frequent on the outer wall and rare on the middle walls.
Remarkably, all Scutellospora spp. lacking a middle wall (morphological
group Cl) have a staining reaction on the outer wall — similar to all species in
Gigaspora but with a different staining intensity from yellow to yellow-orange
to orange-red to red brown — but generally lack a staining reaction on the
inner wall (TABLE 2).
Molecular phylogeny
The 18S and 25S rDNA-based clades generated by the two independent
molecular phylogenetic reconstructions of the family Gigasporaceae were
congruent and indicate polyphyly and paraphyly in the current classification
(PLATES 2-3). Although the alignments show slight differences in tree topologies
and species sequence composition, the major clades are identical. Strikingly,
the clades obtained support our morphological analyses based mainly upon
germination shield characteristics.
The 25S rDNA trees utilized a smaller number of Scutellospora species
sequences than the 18S rDNA trees. Despite this, the fully resolved 25S rDNA
trees comprised all six morphological groups based on examination of spores
from all species described in this genus. For that reason we use PLATE 3 to show
the molecular support for placing monophyletic groups into new families and
genera.
The molecular reconstruction confirmed that Gigaspora derives from within
Scutellospora and not the other way around as hypothesized by Morton (1990)
and Bentivenga & Morton (1995). Here, Scutellospora sequences cluster in three
A, Band C clades (PLATES 2-3). All three clades received high bootstrap values,
regardless of tree-building method used (PLATE 3). The proposed new families
are based on these three clades. The phylogenetic trees further show sub-clades,
which we name at this stage A, B1, B2, B3, Cl, C2 (PLATES 2-3). Although
some sub-clades represent only one or few species sequences, they are further
supported by our morphological analyses.
Group A (formed by S. calospora and related species) does not share the
same most recent common ancestor either with the other two Scutellospora
clades (B and C) or with Gigaspora in the 18S and 25S trees (PLATES 2-3),
indicating polyphyly. However, although the 25S rDNA-generated distance
tree implies paraphyly, it also fully supports monophyly for the groups used
by us to propose new families. All Group A 25S sequences possess an indel
(insertion-deletion sequence) shared only with the outgroup Acaulosporaceae
‘
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New taxa in the Glomeromycetes ... 323
: A AJ276093 S. aurigloba :
| AJ306443 S. calospora Scutellosporaceae !
! AJ306437 S. nodosa fam. nov.
SA eae AJ242729 S. projecturata !
'B 98,96,97 AJ871273 S. reticulata 2 7
| 88,59,50 | AB041345 S. cerradensis Ponhiscutiiirode
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PLATE 2— FIG. 10. Phylogenetic reconstruction of the family Gigasporaceae based on nearly
complete 18S rDNA sequences. The tree was generated using Maximum Likelihood criteria with
1000 bootstrap replicates using the GTR substitution model (Lanave et al.1984) + I + G with the
following parameters: a = 2.0779; b: 10.9922; c: 3.3253; d: 0.7967; e: 18.3736; Nucleotide frequencies:
pi(A) = 0.2740; pi(C) = 0.1857; pi(G) = 0.2541; pi(T) = 0.2862; number of substitutions types 6,
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by capital letters A, B and C, as well as the sub-clades B1, B2, Cl and C2. Nore: sequences of sub-
clade B3 unavailable for these trees.
324 ...Oehl, de Souza & Sieverding
Scutellosporaceae
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100/100/94 EU252109 S. calospora
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Gigasporaceae TUNIS AY 900506 G. gigantea 1
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emend. AF396783 G. margarita :
91/95/89 hae
66/100/75 ¥12075 G. rosea :
Racocetraceae 94/98/80. AY639261 S. pellucida
fam. nov. 87/95/91 AY639323 S. pellucida )
AF396784 S, pellucida
mecha hd Cy AY900508 S. verrucosa
71/68/82
AY900507 S. verrucosa
$1/100/94
if) Y 12076 S. castanea
98/74/98 AJ510232 S. gregaria
AY 900509 S. verrucosa
AY900511 S. verrucosa
AY900510 S. verrucosa
~~ 1() changes
PLATE 3-FIG. 11. Phylogenetic reconstruction of the family Gigasporaceae based on partial 25S
rDNA sequences. The tree was generated using Maximum Likelihood criteria with 1000 bootstfap
replicates using the GTR substitution model (Lanave et al. 1984) + G substitution model with the
following parameters: a = 0.4967; b: 7,3879; c: 2.4411; d: 0.8468; e: 15.7499; Nucleotide frequencies:
pi(A) = 0.3057; pi(C) = 0.1423; pi(G) = 0.2535; pi(T) = 0.2985; Number of substitutions types
6, rates gamma, gamma distribution parameter alpha: 0.5688, proportion of invariable sites = 0.
The tree was rooted with Acaulospora (AY900514, AJ510228) as outgroup. Bootstrap values for
topologies obtained by Maximum Parsimony with gaps treated as missing character; Maximum
—
a
ite,
ee
New taxa in the Glomeromycetes ... 325
(see alignment ALIGN_001257 deposited at EMBL database) indicating a
common evolutionary event. The 18S-based tree could not fully resolve this
clade, as it appears as a polytomy. Group B encompasses species sequences of
three morphological groups based on our analyses. ‘The trees strongly support
separation of Group B2 from B1 and B3 (PLATE 3). Separation between B1 and
B3 is less evident as both seem to descend from the same last common ancestor.
Nevertheless, the remarkable morphological differences between species in B1
and B3 provide full support for our proposed taxonomic changes. Subgroups
C1 and C2 within Group C are clearly separated in both 18S and 25S rRNA
trees.
Our analyses clearly show that Gigasporaceae with its two old genera
Gigaspora and Scutellospora do not reflect the evolutionary history of this
group. The polyphyletic/paraphyletic nature of Scutellospora and the position
of the Gigaspora clade within the Scutellospora clades solicitates the revision
of the Gigasporaceae and supports the erection of three new families based on
the clades A, B and C within the Diversisporales. The molecular phylogenetic
reconstruction also gives some support for the erection of five new genera
within the families erected from clades B and C (PLATE 3). Although the sub-
groups within clade B were represented by sequences of only one or two species
in both trees, the morphological differences are conspicuous (TABLE 2): B1,
represented by S. heterogama; B2, represented by S. nigra, and B3 represented
by S. erythropus. Clade C clearly shows two well supported sub-clades: C1 and
C2 (PLATE 3), with species belonging to each sub-clade possessing significant
morphological spore differences (TABLE 2). Finally, the family Gigasporaceae
remains with a single genus Gigaspora.
Re-organization of Scutellospora based on morphological and molecular
analyses
Based on the morphological differences of germination shields and spore wall
numbers (TABLE 2) and supported by the above phylogenetic analyses, (PLATES
2-3) we can attribute current Scutellospora species to three higher level (A, B, C)
groups and six subgroups (A, B1, B2, B3, C1, C2); Gigaspora species form another
distinct higher level group. Based on the higher groups, we first emend the
_ Gigasporaceae and describe three new families in the Diversisporales. Further,
from within Scutellospora we assign 10 species to genetic group A, 9 to Group
C1, 5 to Group C2, 4 to Group B1, 7 (including one new species see below) to
Group B2, and 1 to Group B3. We hereafter emend the genus description of
_ Parsimony considering gaps as fifth state, and Maximum Likelihood criteria are shown respectively.
The thickest branch lines designate monophyletic groups used to erect new families; the second
_ thickest branch lines show the genera. The three major Scutellospora clades are indicated by capital
letters A, B and C, as well as the sub-clades B1, B2, B3, Cl and C2.
326 ...Oehl, de Souza & Sieverding
Scutellospora, which corresponds to Group A, and describe five new genera —
corresponding to Cl, C2, B1, B2 and B3. Following the International rules, we
select a type species for each new genus from each genetically related group ©
that was first described in literature. One exception was made for S. heterogama
because of the particular taxonomic problems with this species (see below).
Finally, we present identification keys to the families, genera and species with
gigasporoid spore formation and list (TABLE 3-4) the morphological features
obtained from all species in the former Scutellospora under the new generic |
headings.
Taxonomic problems encountered during study
While revising the Gigasporaceae, we encountered some species mis-
identifications and other problems noted below.
Isolates analysed and identified as Scutellospora heterogama by Franke &
Morton (1994) and Jeffries et al. (2007) do not correspond to the S. heterogama
type specimens deposited at FH (as Endogone heterogama). While S. heterogama
sensu Franke & Morton (1994) and Jeffries et al. (2007) has a simple violin-
shaped to oval germ shield, the germ shield of the FH type of E. heterogama is |
ellipsoid and has a dentate periphery and several gti. Also, the type spores are |
lighter in color (pale yellow-brown; Nicolson & Gerdemann 1968) than the
red- to dark-brown spores of S. heterogama sensu Franke & Morton (1994) and |
Jeffries et al. (2007). For these reasons we cite below the type of E. heterogama |
in a new combination and describe S. heterogama sensu Franke & Morton and
Jeffries et al. as a new species in a different genus.
Secondly, our examination of type materials deposited at OSC and FH
revealed that S. weresubiae has only one inner wall, which does not stain in |
Melzer’s, while the outer wall turns yellow to orange-yellow, a finding clearly |
— SSS
different from the original description. We believe that Koske & Walker (1986) |
identified the thin innermost wall layer of the outer wall as a separating middle |
wall (Koske & Walker 1986).
Thirdly, Scutellospora scutata sensu Walker & Diederichs (1989) was
described with three spore walls, while S. scutata sensu Silva et al. (2006b; —
based on INVAM reference accession BR243) has four spore walls. The germ |
shields in both have a dentated periphery in planar view but photographs of the —
germination shield in INVAM'ss S. scutata as published in Walker & Diederichs |
(1989) and here (FIG. 43, photographer F. Oehl) suggest two different species.
We do not believe that the germ shield differences represent just an intra-species |
variation, but in the absence of complete information, we do not describe the —
INVAM specimen as a new species.
Next, we attributed S. trirubiginopa to Group B1 although it was not totally |
clear from the species description that the brown germination shield was indeed |
New taxa in the Glomeromycetes ... 327
bi-lobed. Specimens were not available, and photographs of spores and the
shield were not published. However, the description says that the germination
shield is simple and that two germ tubes might arise from the shield.
For S. crenulata only a few spores were available to us, and the shields, hidden
under the thick and conspicuously ornamented outer spore wall, were difficult
both to detect and to isolate from the rw. Moreover, several infolds prevented
us from determining the number and organization of the lobe(s) (FIGs. 19-20)
and whether more than one gti was present.
Finally, we did not consider a sequence of S. dipapillosa (Z14013) in our
molecular analyses as we have doubts about the correct species identification
— it is likely that there was a confusion of S. dipapillosa with S. heterogama sensu
Franke & Morton (1994) and Jeffries et al. (2007).
Molecular problems encountered during study
In the Glomeromycota the 18S rDNA genes possess low phylogenetical
signal, and trees generated from 18S rDNA sequences did not fully resolve the
ancestry of the clades B, C and Gigaspora, with Clade C receiving the lowest
bootstrap values (PLATE 2), which point to a conflict in the position of clade
C on 18S-based trees. On the other hand, the 25S rDNA-generated trees were
fully resolved; the main clades used to erect new families received bootstrap
values >66% among all tree-building methods used, and >80% for Maximum
Likelihood criteria. Despite the lower number of available species sequences,
‘groups obtained on the 25S rDNA trees are congruent with the germination
shield-based morphological groups.
: CLADE A — At one hand, the 18S rDNA based trees could not resolve the
terminal nodes, which appear as a polytomy. On the other hand, the 25S
rDNA trees were built using only sequences of a single species, S. calospora.
Nonetheless, both tree topologies (PLATES 2-3) agreed with the placement of
clade A (see also de Souza et al. 2005).
CLADE B - The 25S rDNA trees show support for the three proposed genera
on this clade, but they were represented by sequences of only one species for
each genus, while in the 18S rDNA based tree, there is no sequence of group
B3, and differences between B1 and B2 are less resolved, due to the position of
S. cerradensis sequences. The germination shield in S. cerradensis (FIG. 42; see
also S. hawaiiensis, FG. 41) differs to some extent from the typical dentate
shields in D. nigra and all other Dentiscutata species (FIGS. 38-44), although
it clearly resembles more the latter than the bi-lobed shields in Fuscutata
ispecies (PLATE 8). Further work is needed to resolve the relationships of
S. cerradensis and S. hawaiiensis.
CLADE C - Clade C is well resolved on the 25S rDNA trees, although the sub-
clade C1 terminal nodes appear as a polytomy and the S. verrucosa sequences
328 ...Oehl, de Souza & Sieverding
appear non-monophyletic. We intentionally selected those S. verrucosa —
sequences to show this feature of AME The rDNA copies are polymorphic —
in the genome of all Glomeromycota species studied to date (see de Souza et —
al. 2004). Due to this intraspecific rDNA polymorphism, non-monophyly —
among rDNA copies obtained from a single species is common throughout the _
Glomeromycota. However, such polymorphism does not affect the resolution of |
the rDNA-based phylogeny.
Taxonomic revision of Gigasporaceae
Key to families and genera with spores formed on bulbous sporogenous cells
(Diversisporales)
1a Spores with one spore wall; generally with germ warts on inner surface of the wall
Ltn AtULe.S POLES oe ae ee FAMILY 1: Gigasporaceae (Gigaspora) —
1b Spores with outer spore wall and one to three inner walls; germination shield on
Ler MOstiwall hes cee cares elaine Be ee eee CYR ER LARP aa ity 2
2a Spores with a hyaline to subhyaline seldom light yellow germination shield ....... 3}
2b Spores with a yellow brown to brown germination wall
FAMILY. 4: Dentisentalgecde (3/Senel a). ae eee eet a ee 4 ||
3a Germ shield simple, generally (mono-) to bi-lobed, thus (1-)2 germ tube initiations |
(Stiespores Withhoewallsy mes jens on FAMILY 2: Scutellosporaceae (Scutellospora)
3b Germ shield multiple-lobed, 4 or more compartments with each one gti
FA MI Yas: -Racocetraceae Qicenera) seis te hee ee 5 |
4a Germination shield simple, generally bi-lobed; spores with 3 walls ...... Fuscutata |
4b Germination shield complex, >8 compartments, each with a gti; shield periphery
generally dentate in planar view; spores with three or more walls ............. 6
Saxspores with 2 walls eas Sage antl eit oti a Sheree ote Racocetra »
Sb Sporesawith Siwall suet alah a eee a ae et ee Cetraspora |
Ga-Sporesiwithswall ss cjaide feseaeny eee hy bee he Ds a eS Dentiscutata
6b. spores withy>3 Wallse en ies cae ee ne eae glee eee ke Oe ee es Quatunica |
Gigasporaceae J.B. Morton & Benny emend. Sieverd., FA. Souza & Oehl
EMENDATION: Sporocarps are unknown. Spores formed singly in soil or rarely in
roots. Spores are formed on bulbous sporogenous cells arising from subtending
hyphae (sporophore) that differentiate from mycelia hyphae in soil. Spores with |
one spore wall consist of three layers: a unit, semi-persistent to persistent outer ”
layer, a laminate middle layer and a thin inner germinal layer. The germinal |
layer has multiple, randomly or regularly organized germ warts. From one or ©
sometimes several of the germ warts a germ tube arises during germination,
penetrates directly the spore wall and generally branches in a short distance
from the spore. Auxiliary cells formed in hyphal mycelium have spiny or |
nodulous elevations but not smooth round. Form arbuscular mycorrhiza; so
far vesicles in roots unknown.
a ne
—————
New taxa in the Glomeromycetes ... 329
TYPE GENUS: Gigaspora Gerd. & Trappe emend. C. Walker & EE. Sanders,
Mycotaxon 27: 179. 1986.
TYPE SPECIES: Gigaspora gigantea (T.H. Nicolson & Gerd.) Gerd. & Trappe,
Mycologia Memoir No. 5: 29. 1974.
EXCLUDED GENUS: Scutellospora
Key to Gigaspora species
For the key we considered the original descriptions of the species. The synonomy of G. rami-
sporophora with G. margarita (Bentivenga & Morton 1995) is not accepted based on molecular (de
Souza et al. 2004) and morphological (Spain et al. 1989) differences. Gigaspora rosea and G. candida
are also treated as separate species. We exclude G. tuberculata from Gigaspora (see also Bentivenga
~ & Morton 1995).
la. Mean diameter of mature spores < 250 um; spores rarely > 300 um.............. 2
1b. Mean diameter of mature spores > 250 um; spores generally > 300 um........... 3
BeTeneony Tite WUET YOUNG Lehto eee EB are ne ur ema eke ee ee +
2b Spores white to cream, usually with a rose-pink tint; 230-300 um in diam;
sporogenous hyphae darker than the spore wall color ................. G. rosea
3a Spores pearly white when young, 200-490 um in diam.....................045. 5
_ 3b Spores generally pigmented already when young; 290-810 um in diam; mature
spores brilliant yellow-green, color associated with spore contents; spore
contents turning bright red in alkaline solutions; laminae of spore wall not
turning dark purple in Melzer’s reagent; composite wall thickness generally
cel OslitNiaete te eeeyeene thon eter eth, NASA SN PIO NSN lA ene G. gigantea
| 4a. Mature spores dull white with sporogenous hyphae similar in color with
BPOLGAWal Meee ee Meco re eit ree ee a et ctone me Tae Ee ae pe Rae 6
_ 4b. Spores (orange) white to white orange, 140-270 um in diam .... G. alboaurantiaca
| 5a Spores with one sporogenous hyphae; 260-480 um in diam mature spores pearly
white to an opaque, creamy yellow color; composite wall thickness generally
& DES TATETIN A TETANY Oe OS PR noe A reer hoes eee Nee Sey Pea G. margarita
: 5b Spores with composite spore wall thickness generally > 25 um, 200-500 um
iediaim jiature spores yellow (0 DIOWN fee. Pr ty ier ee ee 7
6a Spores with composite spore wall thickness generally < 8 um, 200-300 um in
diam, average 250 um, sporogenous cell dull white 30-50 um diam
«ost Wg otic cnes At Se eee ORAS ara ae YAMA Cer AL AMES oo BP G. candida
| 6b. Spore with composite spore wall thichness < 12 um, 143-350 wm in diam,
average 265 um, sporogenous cell hyaline to yellow 24-50 um diam..... G. albida
| 7a Spores with composite spore wall thickness generally > 25 um, 320-490 um
in diam; mature spores golden yellow to brown, with one sporogenous cell
TE HABA ok: fd em HORM ORE DEAS EPEAT REAR ER NOME L coed Fn SP G. decipiens
_7b. Spores as in 6a, subtending hypha forms one or more sporogenous cells one
on top of other appearing as beads, spore formed on last cell; from any of
the sporogenous cells another subtending hypha can emerge and may or
may not form another sporogenous cell from which another spore forms
er oe ie ee ts Cele atk ae eee G. ramisporophora
330 ...Oehl, de Souza & Sieverding
Scutellosporaceae Sieverd., RA. Souza & Oehl, fam. nov.
MB 511945
Sporae terminaliter efformatae anguste adiacetae ad cellulas sporogeneas; cum tunicis
tribus; scutellum germinale coniunctum ad tunicam interiorem, viola-forme vel ovale
vel cardioforme, hyalinum ad alboflavum, cum 1-2 lobis et depressionibus germationis;
formans structuras mycorrhizarum arbuscularum.
KEY CHARACTERS: Sporocarps are unknown. Spores generally singly formed on |
bulbous sporogenous cells that are formed terminally on a subtending hypha |
that arises from mycelia hyphae in soil. Spores have 3 walls, an outer, a middle
and an inner, germinal wall. Outer wall with 3 or several layers, middle wall
with 1-2 layers, and inner wall with 2 or several layers. A germination shield
is formed on the outer surface or between the outer and the subsequent layer
of the inner wall. Germination shield is transparent, or hyaline to subhyaline,
seldom light yellow, generally bi-lobed, rarely mono-lobed; often violin-shaped
or oval to ovoid, rarely cardioid, circular or coiled; only a few folds cover the
shield surface where in general two germ tube initiations (gti) are formed. |
Generally from one gti, seldom from both gti, a germ tube arises and penetrates _
the outer spore walls. Subtending hyphae form one to several septa in some |
distance to the sporogenous cells. Auxiliary cells in the hyphal mycelium are, as
far as known, knobby without spines on the surface. Forming typical arbuscular "|
mycorrhizae; vesicle formation in roots is so far unknown.
TYPE GENUS: Scutellospora C. Walker & EE. Sanders emend. Oehl et al.
Scutellospora C. Walker & EE. Sanders. emend. Oehl, F.A. Souza & Sieverd.
(
EMENDATION: Sporocarps unknown. Spores formed on sporogenous cells that |
form terminally on a hypha which arises from mycelia hyphae in soil. Outer
spore wall generally is three-layered and continuous with the wall of the |
sporogenous cell. Outer layer of the outer spore wall generally rigid, second |
layer laminate and third layer thin, often membranous, tightly adherent to
the laminate layer and thus, often difficult to observe. Pore between the spore
and sporogenous cell is narrow and usually closed by a plug formed by spore :
wall material. Two hyaline walls (Mw’ and ‘Iw’) form de novo during spore |
formation and have 1-2 and 2-3 layers, respectively. The tw is two to three-
layered forming a germination shield on its outer surface or between the outer |
and the subsequent layer of rw. Germination shield is transparent, or hyaline |
to subhyaline, seldom light yellow, bi- to mono-lobed; often violin-shaped |
to oval to ovoid to more rarely cardioids or coiled and then, either circular’
or apparently broad ellipsoid to irregular; only a few folds cover the shield —
surface where 1-2 rounded germ tube initiations (‘gti, about 2-4 um in diam) |
ae
New taxa in the Glomeromycetes ... 331
are visible from where the germ tubes arise which penetrate the outer spore
wall layers. Mycelia hyphae form one to several septa in some distance to the
sporogenous cells. Auxiliary cells in the hyphal mycelium, as far as known, are
knobby without spines on the surface. Forming typical arbuscular mycorrhizae,
vesicle formation in roots so far unknown.
TyPE SPECIES: Scutellospora calospora (T.H. Nicolson & Gerd.) C. Walker &
FE. Sanders
Key to Scutellospora species
Sepiiores iL OULOfnammentatiomon outer, Walia) verte lene ae pic cece electron epee 2
Sei iLL OT amentalion, OMOuteh Wall ay. 24 icc tie es ls cow's Ale cies Vas as oe 3
2a Spores light colored: subhyaline, pink, creamy, straw to greenish or
Bowalishiy cll Wiehe went ei ree 00/55 '2h oe ag Peete ase ed east 4
Beerecyy CHOW. Drown (0) DIAC ammeter gs. 5 2oie ee vth oes oce s oe 5
3a Spores with one homogenous ornamentation on spore surface .................. 6
Sp spores with two-type ornamentation on spore surface ...,...........--++++:0:- 7
4a Spores subhyaline to creamy to pale straw to greenish yellow.................... 8
Ab Spores dark yellow to golden yellow to brown yellow ....................-.005 9
5a Spores yellow brown to orange-brown, 160-360 um in diam, outer and
innermost wall strongly staining in Melzer’s reagent ............... S. arenicola
5b Spores dark brown to black, 300-460 um in diam ................... S. tricalypta
6a Spores hyaline to pale yellow, 160-270 um in diam; with knobs 3.5-6.5 um
Dae nrandas Oo O,oaim Widerat baseauc sine nee bak). os eas ee S. nodosa
6b Spores golden yellow to ochraeous to sienna, 100-180 um in diam; with
2.0-4.0 um long protuberances formed by the structural, laminated layer
: SM CLOULCI WAL OMe ete Mian ty ene eter. «aetna S. projecturata
| 7a Spores with two-type ornamentation on spore surface; spores pale orange
brown to dark orange brown, 130-180 um in diam; with small conical
warts and additional blunt, bacilliform larger projections on the spore
| MUTE on SO Es ok ae ee a PRR AIS Som are S. dipapillosa
7b Spores with warted projections having central secondary projections in the tip;
Bares cceaintovellow, 100-170 umn diam a yom... 55 ee S. crenulata
8a Spores subhyaline to pale straw to pale greenish-yellow, (100—)150-300(—500) um
in diam, generally ellipsoid to oblang, with bi-layered middle wall .. .S. calospora
8b Spores yellow to greenish-yellow; 140-240 um in diam, generally globose to
subglobose, with one layered middle wall.................... S. dipurpurescens
9a Spores golden yellow to dull yellow, (130-)200-420(-520) um ........ S. aurigloba
9b Spores dark yellow to brown yellow (to yellow brown), 105-150 um, outer spore
wall strongly expanding in lactic acid based mountants........ S. pernambucana
332 ...Oehl, de Souza & Sieverding
PLATE 4—Typical simple, hyaline to subhyaline, violin-shaped to oval, to rarely cardioid, circular, |
oblong or irregular germination shields in Scutellospora spores. Spores have one outer (ow), one |
middle (mw) and one inner (1w) spore wall. Generally, shields have two compartments (lobes) —
with one germ tube initiation (gti) per lobe from where the germ tubes (gt) may arise (germination | |
visible in F1G. 12). Rarely shields are mono-lobed (FIGs. 19-21). Shield and inner wall in S. tricalypta
darkened during 25 years mounted in Ferrant medium (FIG. 14).
a
New taxa in the Glomeromycetes ... 333
Scutellospora arenicola Koske & Halvorson,
Mycologia 81: 927. 1990 [‘1989”]. (FIG. 15)
Scutellospora aurigloba (1.R. Hall) C. Walker & RE. Sanders.,
Mycotaxon 27: 180. 1986; emend. C. Walker & LR. Hall,
Mycol. Res. 95: 400. 1991. (FIG. 17)
BASIONYM: Gigaspora aurigloba LR. Hall, Trans. Brit. Mycol. Soc. 68: 351. 1977.
Scutellospora calospora (T.H. Nicolson & Gerd.) C. Walker & EE. Sanders,
Mycotaxon 27: 180. 1986. (FIG. 12)
BasionyM: Endogone calospora T.H. Nicolson & Gerd., Mycologia 60: 322. 1968.
= Gigaspora calospora (T.H. Nicolson & Gerd.) Gerd. & Trappe,
Mycologia Memoir No. 5: 28. 1974.
Scutellospora crenulata R.A. Herrera, Cuenca & C. Walker,
Can. J. Bot. 79: 674. 2001. (FIGS. 19-20)
Scutellospora dipapillosa (C. Walker & Koske) C. Walker & EE. Sanders,
Mycotaxon 27, 181. 1986. Gare, 18))
BASsIONYM: Gigaspora dipapillosa C. Walker & Koske, Mycologia 77: 709. 1985.
Scutellospora dipurpurescens J.B. Morton & Koske,
Mycologia 80: 520. 1988. (FIG. 16)
Scutellospora nodosa Btaszk., Mycologia 83, 537. 1991. (FIG. 18)
Scutellospora pernambucana Oehl, D.K. Silva, N. Freitas, L.C. Maia,
Mycotaxon 106: 363. 2008. (FIG. 21)
Scutellospora projecturata Kramad. & C. Walker, Annals Bot. 86: 22. 2000.
Scutellospora tricalypta (R.A. Herrera & Ferrer) C. Walker & FE. Sanders,
Mycotaxon 27: 180. 1986. (FIG. 14)
BASIONYM: Gigaspora tricalypta R.A. Herrera & Ferrer, Rev. Jard. Bot. Nacional, Habana. 1:
49. 1981 [‘1980’].
-Racocetraceae Oehl, Sieverd. & EA. Souza, fam. nov.
MB 511946
Sporae terminaliter efformatae anguste adiacetae ad cellulas sporogeneas; scutellum
germinale coniunctum ad tunicam interiorem, hyalinum subhyalinum ad alboflavum,
undulo-lobatum cum 4-12 lobis et 4-12 depressionibus circularibus germinationis;
formans structuras mycorrhizarum arbuscularum.
_ FIG. 12. Type species S. calospora (germination visible). FIG. 13. S. dipapillosa F1G. 14. S. tricalypta
(25-year old spores in Ferrant medium). FIG. 15. S. arenicola. FIG. 16. S. dipurpurescens. FIG. 17.
: S. aurigloba. F1G. 18. S. nodosa. FIGS. 19-20. S. crenulata (monolobed). Fic. 21. S. pernambucana
-(monolobed).
334 ...Oehl, de Souza & Sieverding
KEY CHARACTERS: Spores formed singly in soils and rarely in roots formed on
bulbous sporogenous cells which are formed terminally on a subtending hypha
that arises from mycelia hyphae. Spores with two to three walls: an outer spore
wall, either no middle wall or one middle wall, and an inner germinal wall.
Each wall if present has one, two or several layers. A germination shield arises
on the outer surface or beneath a thin outer layer of the inner wall; germination
shield is hyaline to subhyaline seldom light yellow, generally oval to ellipsoid
or subglobose, with several (4-12) conspicuously lobed, wave-like projections
forming the outer surface of the shield; folds separate the lobes in compartments
on the shield, and each compartment may have a germ tube initiation (gti).
From one or rarely two gti a germ tube may arise and penetrate the middle
and the outer spore wall. In the substending hypha of the sporogenous cell,
often several septae are formed in some distance from pore. Auxiliary cells in
the hyphal mycelium as far as they are known knobby without spines on the
surface. Forming typical arbuscular mycorrhizae; vesicle formation in roots
unknown. |
TYPE GENUS: Racocetra Oehl et al.
OTHER GENUS: Cetraspora
Racocetra Oehl, EA. Souza & Sieverd., gen. nov.
MB 511947
Sporae terminaliter efformatae anguste adiacetae ad cellulas sporogeneas; cum tunicis
duabus; scutellum germinale coniunctum ad tunicam interiorem, hyalinum ad alboflavum,
undulo-lobatum, 4-12 lobis et 4-12 depressionibus germinatione; formans structuras
mycorrhizarum arbuscularum.
ETtyMoLoGcy: from the Greek: (paxoc, racos = cloth, lobe), and from the Latin: cetra
(light shield), referring to the wavy-lobed surface of the germination shield in planar
view.
KEY CHARACTERS: Spores formed singly in soil and rarely in roots, on bulbous
sporogenous cells that arise terminally on mycelia hyphae. Outer spore wall is
generally three-layered and continuous with the wall of the sporogenous cell.
Outer layer of the outer spore wall generally semi-persistent to persistent, rigid;
second layer laminate; third layer thin, often membranous, tightly adherent to
the laminate layer and thus, often difficult to observe. Pore between spore and
sporogenous cell is narrow and usually closed by a plug formed by spore wall
material. A single inner wall forms de novo during spore formation and has
two to three layers. A germination shield arises on the outer surface or beneath
a thin outer layer of the inner wall; germination shield is hyaline to subhyaline
seldom light yellow, generally oval to ellipsoid or subglobose, with several (4-
12) wave-like lobed projections forming the outer surface of the shield; folds
separate the lobes on the shield, and each lobe may have a germ tube initiation
New taxa in the Glomeromycetes ... 335
PLATE 5—Racocetra germination shields (gs) have multiple compartments that give a multiple-
lobed, wavy shield appearance. Racocetra spores have one outer (OW) and one inner (Iw) spore
wall. Generally one germ hole (gh) present with one germ tube initiation (gti) visible on each of
the multiple lobes.
Fic. 22-23. Type species R. coralloidea. Fic. 23. Germ shield wall in the type specimen
partly darkened after more than 30 years of storage in lactophenol. Fic. 24. R. gregaria. Fic. 25.
R. castanea. Fic. 26. R. persica. Fic. 27. R. verrucosa. Fics. 28-29. R. fulgida. Fic. 29. Shield in
a newly developed spore; gti’s not yet formed. Fics. 30-31. R. weresubiae shield not completely
visible in planar view and thus overlapping itself; slightly deformed through spore crushing;
Fic. 31. Shield wall amplified using Adobe® Photoshop 9.0.
(about 2-5 um in diam) from where the germ tubes arise and penetrate the
outer wall. In the subtending hypha of the sporogenous cell one to several septa
formed in some distance to the sporogenous cells. Auxiliary cells in the hyphal
mycelium (as far as they are known) knobby without spines on the surface.
Forming typical arbuscular mycorrhizae; vesicle formation in roots is so far
unknown.
TyPE SPECIES: Racocetra coralloidea (Trappe, Gerd. & I. Ho) Oehl et al.
336 ...Oehl, de Souza & Sieverding
Key to Racocetra species
la Spores without ornamentation.on Outer wallWe weiss tact een ee 2
Ib Spores with ornamentation on outerawalll se gece a. eee eee 3
2a Spores light cOlored fries ce) elves: pew chs shone cle the ene eet eee 4
2b Spores creamy-brown to brown-sienna, 170-370 um in diam.......... R. castanea
3a Surface ornamentation with spines or warts generally <2.5umindiam.......... 5
3b Surface ornamentation with projections generally > 2.5 um in diam ............. 6 |
4a Spores hyaline to pale straw to pale yellowish-cream, 160-250 tm in diam
Ee ee ees red i eer Co cane Ports MO tes ie tr ....R. fulgida ||
4b Spores pale to dark pink, 120-170 x 130-420 um in diam; outer wall staining
yellow to yellow orange in Melzer’s reagent ........ (R. alborosea)/R. weresubiae
5a Spore surface with fine spines; spores pinkish-orange to brownish orange,
2707390 Wma Deiat, 00 Joy ise eree een eee eee R. persica
5b Spore surface with crowded rounded warts; spores pale straw to yellow to light
orange brow 20-480) (mi ciate eee R. verrucosa |
6a Spores red brown to dark brown with rounded projections on surface............ 7
6b Spores dark brown, 300-460 um in diam; with coralloidic warts...... R. coralloidea |
7a Spores red brown to dark brown, 250-450 um in diam; with rounded
dome-shaped wWartsiyst. we teens choi fr ot ee rye R. gregaria |
7b Spores brown to dark brown; 90-180 um in diam; warts on top with
centralidepressions. # acuta «cries tees Sens ee a eee R. minuta |
Racocetra alborosea (Ferrer & R.A. Herrera) Oehl, EA. Souza & Sieverd., comb. nov.
MB 511950
BASIONYM: Gigaspora alborosea Ferrer & R.A. Herrera, Rev. Jardin Bot. Nacional,
Habana. 1: 55. 1981 [‘1980’].
= Scutellospora alborosea (Ferrer & R.A. Herrera) C. Walker & FE. Sanders,
Mycotaxon 27: 180. 1986.
Racocetra castanea (C. Walker) Oehl, RA. Souza & Sieverd., comb. nov.
MB 511952 (FIG. 25)
BastonyM: Scutellospora castanea C. Walker, Cryptogamie Mycologie 14: 280. 1993.
Racocetra coralloidea (Trappe, Gerd. & I. Ho) Oehl, R.A. Souza & Sieverd.,
comb. nov. (FIGS. 22-23)
MB 511948
BasIONYM: Gigaspora coralloidea Trappe, Gerd. & I. Ho, Mycologia Memoir No. 5:
30. 1974.
= Scutellospora coralloidea (Trappe, Gerd. & I. Ho) C. Walker & EE. Sanders,
Mycotaxon 27: 181. 1986.
Racocetra fulgida (Koske & C. Walker) Oehl, EA. Souza & Sieverd.,
comb. nov. (FIGS. 28-29)
MB 511955
BastonyM: Scutellospora fulgida Koske & C. Walker, Mycotaxon 27: 221. 1986.
New taxa in the Glomeromycetes ... 337
Racocetra gregaria (N.C. Schenck & T.H. Nicolson) Oehl, F.A. Souza & Sieverd.,
comb. nov. (FIG. 24)
MB 511949
BASIONYM: Gigaspora gregaria N.C. Schenck & T.H. Nicolson, Mycologia 71: 185.
1O79)
= Scutellospora gregaria (N.C. Schenck & T.H. Nicolson) C. Walker & EE.
Sanders, Mycotaxon 27: 181. 1986.
Racocetra minuta (Ferrer & R.A. Herrera) Oehl, EA. Souza & Sieverd., comb. nov.
MB 511951
BASIONYM: Gigaspora minuta Ferrer & R.A. Herrera, Rev. Jardin Bot. Nacional,
Habana. 1: 53. 1981 [‘1980’].
= Scutellospora minuta (Ferrer & R.A. Herrera) C. Walker & EE. Sanders,
Mycotaxon 27: 180. 1986.
Racocetra persica (Koske & C. Walker) Oehl, F.A. Souza & Sieverd., comb. nov.
MB 511953 (FIG. 26)
BASIONYM: Gigaspora persica Koske & C. Walker, Mycologia 77: 708. 1985.
= Scutellospora persica (Koske & C. Walker) C. Walker & EE. Sanders,
Mycotaxon 27:181. 1986.
Racocetra verrucosa (Koske & C. Walker) Oehl, EA. Souza & Sieverd., comb. nov.
— -MB511954 (FIGs2 7)
BASIONYM: Gigaspora verrucosa Koske & C. Walker, Mycologia 77: 705. 1985.
= Scutellospora verrucosa (Koske & C. Walker) C. Walker & EE. Sanders,
Mycotaxon 27: 181. 1986.
_ Racocetra weresubiae (Koske & C. Walker) Oehl, EA. Souza & Sieverd., comb. nov.
MB 511956 (FIGS. 30-31)
BASIONYM: Scutellospora weresubiae Koske & C. Walker, Mycotaxon 27: 224. 1986.
Cetraspora Oehl, F.A. Souza & Sieverd., gen. nov.
| MB 511957
Sporae terminaliter efformatae anguste adiacetae ad cellulas sporogeneas; cum tunicis
tribus: scutellum germinale coniunctum ad tunicam interiorem, hyalinum ad alboflavum,
undulo-lobatum, cum 4-12 lobis et 4-12 depressionibus germinatione; formans structuras
mycorrhizarum arbuscularum.
EryMo_oey: from the Latin: cetra (= light shield) and spora (= spore), referring to the
light colored germination shield which often is difficult to observe, as in C. gilmorei and
especially in C. pellucida.
_ Key CHARACTERS — Spores formed singly in soil and rarely in roots on
_ bulbous sporogenous cells that arise terminally on a subtending hypha which
is connected to mycelium. Outer spore wall generally is three-layered and
- continuous with the wall of the sporogenous cell. Outer layer of the outer spore
wall generally semi-persistent to persistent, rigid; second layer laminate; third
layer thin, often membranous, tightly adherent to the laminate layer and thus,
often difficult to observe. Pore between spore and sporogenous cell is narrow
338 ...Oehl, de Souza & Sieverding
and usually closed by a plug formed by spore wall material. Two hyaline walls
(‘middle wall’ and ‘inner wall’) form de novo during spore formation and have
1-2 and 2-3 layers, respectively. A germination shield arises on the outer surface
of the inner wall or beneath a thin outer layer of the innermost wall; germination
shield is hyaline to subhyaline seldom light yellow, generally oval to ellipsoid or
subglobose, with several (4-12) wave-like lobed projections forming the outer
surface of the shield; large folds separate the lobes on the shield, and each lobe
may have an germ tube initiation (about 2-5 um in diam) from where the germ
tubes arise and penetrate the overlaying spore walls. Subtending hyphae form
one to several septa in some distance to the sporogenous cells. Auxiliary cells
in the hyphal mycelium as far as they are known knobby without spines on the
surface. Forming typical arbuscular mycorrhizae; vesicle formation in roots is
unknown.
TYPE SPECIES: Cetraspora gilmorei (Trappe & Gerd.) Oehl et al.
Key to Cetraspora species
la. Spores without ornamentation on outerwall?.. a= see ene y..
lb. Spores:with ornamentation/on outer wall vga) eee 3
2a Spores hyaline to whitewer 7.5 ye Pe eee en ee ~
2b Spores apricot yellow to yellow brown, 140-240 um in diam......... S. armeniaca
3a Spores with a fine dense spiny ornamentation on outer spore wall; spores
ochraeous to fulvous or rust, 130-230 um in diam.............. C. spinosissima
3b Spores ornamentated with finger-print-like processes in planar view; spores
ochraeous yellow with a pinkish tint, 110-190 um in diam............ C. striata
4a Spores brilliant hyaline, white to rarely pale grey, generally globose to
subglobose; (60—)120-—250(—420) um in diam; sporogenous cell a hyaline
to subhyaline; germination shield generally not seen.............. C. pellucida
4b Spores hyaline, becoming creamy under storage in formalin; globose to
subglobose to ellipsoid; 200-320 um in diam; sporogenous cell brown;
germination shield generally easy wo ODSerVenen ss. a ten C. gilmorei
Cetraspora armeniaca (Blaszk.) Oehl, FA. Souza & Sieverd., comb. nov. (FIG. 35)
MB 511961
BAsIONnYM: Scutellospora armeniaca Btaszk., Mycologia 84: 939. 1993.
Cetraspora gilmorei (Trappe & Gerd.) Oehl, EA. Souza & Sieverd., comb. nov.
MB 511958 (FIGS. 32=348
BasIONYM: Gigaspora gilmorei Trappe & Gerd., Mycologia Memoir No. 5: 27. 1974.
= Scutellospora gilmorei (Trappe & Gerd.) C. Walker & EE. Sanders, Mycotaxon
277181.1986.
Cetraspora pellucida (T.H. Nicolson & N.C. Schenck) Oehl, RA. Souza & Sieverd.,
comb. nov. (FIG. 34)
MB 511959
New taxa in the Glomeromycetes ... 339
PLATE 6—Multiple-lobed, wavy germination shield (gs) form in Cetraspora spores; spores with one
outer (ow), one middle (mw) and one inner (Iw) spore wall. On each of the lobes (compartments)
a germ tube initiation (gti) might be formed from where the germ tubes arise. Shields and gti are
generally light or transparent and less obvious than in Racocetra.
FIG. 32-33. Type specimen of C. gilmorei. Spore walls and shield have darkened after > 30 years
stored in lactophenol. Fic. 34. C. pellucida. Fic. 35. C. armeniaca. FIGs. 36-37. C. spinosissima.
BasionyM: Gigaspora pellucida T.H. Nicolson & N.C. Schenck, Mycologia 71: 189.
{979,
= Scutellospora pellucida (T.H. Nicolson & N.C. Schenck) C. Walker & EE.
Sanders, Mycotaxon 27: 181. 1986.
340 ...Oehl, de Souza & Sieverding
Cetraspora spinosissima (C. Walker & Cuenca) Oehl, EA. Souza & Sieverd.,
comb. nov. (FIGS. 36-37)
MB 511960
BASIONYM: Scutellospora spinosissima C. Walker & Cuenca, Annals Bot. 82: 723. 1998.
Cetraspora striata (Cuenca & R.A. Herrera) Oehl, EA. Souza & Sieverd., comb. nov.
MB 512598
BasionyM: Scutellospora striata Cuenca & R.A. Herrera, Mycotaxon 105: 81. 2008.
Dentiscutataceae F.A. Souza, Oehl & Sieverd., fam. nov.
MB 511962
Sporae terminaliter efformatae anguste adiacetae ad cellulas sporogeneas; cum tunicis
tribus vel quartuoribus; scutellum germinale coniunctum ad tunicam interiorem, flavo-
brunneum ad brunneum, viola-forme vel ovale vel cardioforme vel ellipsoideum. Formans
structuras mycorrhizarum arbuscularum.
KEY CHARACTERS: Sporocarps are unknown. Spores formed singly on bulbous
sporogenous cells which are formed terminally on a subtending hypha that
arise from mycelia hyphae. Outer spore wall three- to four to rarely five-
layered. Spores have 3-4 walls, an outer spore wall with several layers, one to
several middle walls and an inner wall each of two or more layers. Germination
shield generally formed on the outer surface of the innermost wall or beneath a
thin outer layer of the inner wall, yellow-brown to brown, violin-shaped to oval
to ovoid to heart-shaped to reniforme to ellipsoid, consisting of two lobe-like
compartments or up to 30 small compartments. Lobes and compartments are
separated by folds and generally have each one germ tube initiations; from one
to a few gtia germ tube may arise penetrating the other walls. Subtending hypha
forms one to several septa in some distance to the sporogenous cells. Auxiliary
cells in the mycelium hyphae as far as they are known knobby without spines
on the surface. Forming typical arbuscular mycorrhizae, vesicle formation in
roots unknown.
TyPE GENUS: Dentiscutata Sieverd. et al.
OTHER GENERA: Fuscutata, Quatunica
Dentiscutata Sieverd., FA. Souza & Oehl, gen. nov.
MB 511968
Sporae singulatim terminaliter efformatae anguste adiacetae ad cellulas sporogeneas; cum
tunicis tribus; scutellum germinale coniunctum ad tunicam interiorem, flavo-brunneum
ad brunneum; cum 8-30 compartimentibus et depressionibus germinationis; formans
structuras mycorrhizarum arbuscularum.
ETyMOLOGy: From the Latin denti(culata) (= dentate), and scutata (= ‘shielded’),
referring to the dentate form of the periphery of the brown germination shield.
KEY CHARACTERS: Sporocarps are unknown. Spores singly formed on bulbous
sporogenous cells that are formed terminally on subtending hypha that arises
New taxa in the Glomeromycetes ... 341
from mycelia hyphae. Outer spore wall is three- to five-layered and continuous
with the wall of the sporogenous cell. Pore between the spore and sporogenous
cell is narrow and usually closed by a plug formed by outer spore wall material.
A hyaline middle wall and a hyaline inner wall form de novo during spore
formation; middle wall one- to bilayered; inner wall two to three-layered.
Germination shield generally arising on the outer surface of the innerwall or
beneath a thin outer layer of the inner wall, yellow-brown to brown, generally
ellipsoid (to oval), or rarely reniforme to cardioforme, usually with many (8-30)
large folds separating the shield into 8-30 ‘small compartments’; each small
compartment generally with one circular germ tube initiation; usually from
one or a few gti germ tubes may arise and penetrate the middle wall and the
outer wall; the periphery of the germination shield generally appears dentate
in planar view. Septa often formed in subtending hypha in some distance to
the sporogenous cells. Auxiliary cells formed in hyphal mycelium as far as they
are known knobby without spines on the surface. Forming typical arbuscular
mycorrhizae, vesicle formation in roots so far unknown.
TYPE SPECIES: Dentiscutata nigra (J.F. Redhead) Sieverd. et al.
Key to Dentiscutata species
Be ooreoviLNOUl Ol namentallon On OULEr Walliams ee meee eet eran nes 2
ie pocesmuli: ornamentation on outer walls. we wear ewe. eels ean 3
2a Spores hyaline, white to light olive, 350-750 um in diam
Sn at Ne A ag Pee Oe D. scutata (sensu Walker & Diederichs 1989)
2b Spores pale orange brown to red brown, 200-300 um in diam; outer wall not
staining in Melzer’s reagent; second middle wall layer fracturing with a series
Sirectangulamandey-shaped NOL es sme a ey ne eee D. hawaiiensis
Eanpores with a single ornamentation on outer spore wall ..........44.......2.5.- 4
Bp opores with double ornamentation on outer spore wall.........2..............- 5
4a Spores hyaline to white, globose to ovoid, 170-370 tm in diam; with spiny
Papillac ornalnentatiOleaes ei. er ene eee ernen eeere gs D. cerradensis
4b Spore with papillae surface ornamentation on outer wall; Spores light brown,
BlODOSeLO cliipsOid@ oO S2OU;IN in Cialis ee, wee een ae D. heterogama
5a Spores with double ornamentation on the outer spore surface.................. 6
5a Spores with each one papillae ornamentation both on outer and inner surface
of the outer wall; yellow-brown to brown, (120—)260-450(-500) um
KWo UPR a) sears: Aeneas Ree ee aes oe Rear Oe) enh peor nen Ae D. biornata
6a Spores orange brown to dark red brown, 200-470 tm in diam; having spines in the
DeDLcssions OL asreLiCUlUIN sation ero, ott lee D. reticulata
6b Spores dark brown to black, 290-500(-1050); with rounded pits and a sinuous
Secondary ornamentation belowseeteitisuetas lanlas aE tioeEy er aeh enna D. nigra
342 ...Oehl, de Souza & Sieverding
Dentiscutata biornata (Spain, Sieverd. & S. Toro) Sieverd., RA. Souza & Oehl,
comb. nov. (FIG. 40)
MB 511971
BastonyM: Scutellospora biornata Spain, Sieverd & S. Toro, Mycotaxon 35: 220. 1989.
Dentiscutata cerradensis (Spain & J. Miranda) Sieverd., RA. Souza & Oehl, comb.
nov. (FIG. 42)
MB 511973
BAsIony™M: Scutellospora cerradensis Spain & J. Miranda, Mycotaxon 60: 130. 1996.
Dentiscutata hawaiiensis (Koske & Gemma) Sieverd., FA. Souza & Oehl,
comb. nov. (FIG. 41)
MB 511972
BastonyM: Scutellospora hawaiiensis Koske & Gemma, Mycologia 87: 678. 1995.
Dentiscutata heterogama (T.H. Nicolson & Gerd.) Sieverd., RA. Souza & Oehl,
comb. nov. (FIG. 44)
MB 511975
BastonyM: Endogone heterogama T.H. Nicolson & Gerd., Mycologia 60: 319. 1968.
= Gigaspora heterogama (T.H. Nicolson & Gerd.) Gerd. & Trappe, Mycologia
Memoir No. 5: 31. 1974.
= Scutellospora heterogama (‘T.H. Nicolson & Gerd.) C. Walker & EE. Sanders,
Mycotaxon 27: 180. 1986.
Dentiscutata nigra (J.F. Redhead) Sieverd., EA. Souza & Oehl, comb. nov.
MB 511969 (FIG. 38)
BASIONYM: Gigaspora nigra J.F. Redhead, Mycologia 71: 187. 1979.
= Scutellospora nigra (J.F. Redhead) C. Walker & EE. Sanders, Mycotaxon 27:
181. 1986.
Dentiscutata reticulata (Koske, D.D. Mill. & C. Walker) Sieverd., EA. Souza &
Oehl, comb. nov. (FIG. 39)
MB 511970
BASIONYM: Gigaspora reticulata Koske, D.D. Mill. & C. Walker, Mycotaxon 16: 429.
1933.
= Scutellospora reticulata (Koske, D.D. Mill. & C. Walker) C. Walker & EE.
Sanders, Mycotaxon 27: 181. 1986.
Dentiscutata scutata (C. Walker & Dieder.) Sieverd., FA. Souza & Oehl, comb. nov.
MB 511974 (FIG. 43)
BastonyM: Scutellospora scutata C. Walker & Dieder., Mycotaxon 35: 357. 1989.
Fuscutata Oehl, EA. Souza & Sieverd., gen. nov.
MB 511963
Sporae terminaliter efformatae anguste adiacetae ad cellulas sporogeneas; cum tunicis
tribus; scutellum germinale coniunctum ad tunicam interiorem, flavo-brunneum ad
brunneum; ovale vel ovoidum vel viola-forme, cum 2 lobis et depressionibus germinatione;
formans structuras mycorrhizarum arbuscularum.
New taxa in the Glomeromycetes ... 343
PLATE 7—Dentiscutata with complex structured, yellow-brown to brown germination shields (gs)
showing several to many (8-30) compartments, separated by large folds (f); often one initial germ
hole (gh) and several to many (8-30) germ tubes initiations (gti) visible; germ tubes emerge from
one or two gti during germination; shield periphery conspicuously dentate in planar view.
) Fic. 38. Type species D. nigra. Fic. 39. D. reticulata. Fic. 40. D. biornata. Fic. 41. D. hawaiiensis
Fic. 42. D. cerradensis. Fic. 43. D. scutata sensu Silva et al. (2006b). Fic. 44. D. heterogama.
EryMo.ocy: from the Latin fu(scus) (= brown), and scutata (= ‘shielded’), referring to
the color of the germination shield formed on the inner spore wall.
KEY CHARACTERS: Sporocarps are unknown. Single spores formed on bulbous
sporogenous cells which are terminally formed on the subtending hypha that
344 ...Oehl, de Souza & Sieverding
arises from mycelia hyphae. Outer spore wall three- to four layered. Pore
between the spore and sporogenous cell is narrow and usually closed by a plug
formed by spore wall material. Two hyaline walls (‘middle wall and ‘inner
wall’) form de novo during spore formation and have 1-2 and 2-3 layers,
respectively. Germination shield generally formed on the outer surface of the
innermost wall or beneath a thin outer layer of the inner wall, yellow-brown
to brown, generally ovoid to violin-shaped to heart-shaped, consisting of 2
lobes and folds; both lobes with a germ tube initiation (gti, about 3-6 um in
diam). Spore germination generally from one gti; germ tube penetrates from
there the middle and outer spore wall. In the subtending hypha one to several
septa are formed in some distance to the sporogenous cell. Auxiliary cells in
the hyphal mycelium as far as they are known knobby without spines on the
surface. Forming typical arbuscular mycorrhizae, vesicle formation in roots so
far unknown.
TYPE SPECIES: Fuscutata heterogama Oehl et al.
Key to Fuscutata species
la Spore without surface ornamentation on outer wall.................... 2222 eee 2
1b Spore with surface ornamentation on outer wall). 224)... 4 ee 5
2a Spores hyaline to white; oblong-ellipsoid to long ellipsoid, 280-580 x
210-370 pumyin diame ee eee Se onan Se EF. savannicola
2b Spores dark orange brown to dark red brown, 140-220 um in diam,
outer and innermost wall strongly staining in Melzer’s reagent.......... E rubra
3a Spores hyaline to subyaline, globose to oval, um in diam; spores with fine
spiny ornamentation on the surface of the outer wall........... FE trirubiginopa
3b Spore with papillae ornamentation on the surface of the outer wall; spores
red brown to dark brown, globose to oval, 159-295 um in diam..... E heterogama
Fuscutata heterogama Oehl, EA. Souza, L.C. Maia & Sieverd., sp. nov.
MB 511964 (FIGS. 45-46)
Sporae singillatim in solo efformatae anguste adiacetae ad cellulas sporogeneas subterminales
vel intercalares, brunneae vel rubro-brunneae vel oscuro-brunneae, globosae, 165-280
um in diametro, vel subglobosae vel ovale (159-265 x 195-295 um); sporae cum tunicis
tribus: tunica exterior stratis tribus in totum 7.0-11.5 um crassa, coniuncta tunicam
cellulae sporogeneae et tunica hyphae; stratum exterius tunicae exterioris hyalinum,
evanescens, 1.1-2.1 um crassum; papillibus ornatum altis 1.1-2.1 um et 0.9-1.5(-2.5) um
latis ornatum; stratum secundum brunneum vel rubro-brunneum vel oscuro-brunneum,
laminatum, 5.5-8.5um crassum; stratum interius tunicae exterioris brunneum vel rubro-
brunneum, subtile; tunica media et tunica interior de novo formans; cum stratis hyalinibus;
tunica media duobus stratis in totum 2.1-2.8 um crassum; tunica interior tribus stratis,
2.1-3.4 um crassa; stratum medium et stratum interiore tunicae exterioris purpureo-
brunneum vel oscuro-brunneum colorans, stratum medium tunicae interioris purpureo
colorans reagente Melzeri; scutellum germinale coniunctum ad tunicam interiorem, flavo-
New taxa in the Glomeromycetes ... 345
eetros =
PLaTE 8—Fuscutata with yellow-brown to brown germination shields and three walls (ow, Mw
and Iw). . Germination shields are generally bi-lobed with a germ hole (gh) and two germ tube
initiations (gti). FIGS. 45-46. Type species E heterogama. Fic. 47. E rubra. ric. 48. Old shield, in
EF, savannicola.
PLATE 9—FIGS. 49-51: Quatunica with one species, Q. erythropus: Complex structured, yellow-
brown to brown germination shields (gs) in spores of type species Q. erythropus having four spore
walls (Ow, MW1, MW2 AND Iw). Germination shield on the outer surface of rw. Shields with many
(> 10) compartments, separated by large folds (f); generally one initial germ hole (gh) and several
germ tubes initiations (gti) visible from where one or few germ tubes per spore emerge during
germination (FIG. 51); periphery of shield walls conspicuously dentate.
346 ...Oehl, de Souza & Sieverding
brunneum ad brunneum; ovale vel ovoid vel viola-forme, simplex bilobatumque cum 2
depressionibus germinationis; structurae mycorrhizarum arbuscularum formans.
TYPE SPECIMENS: BRAZIL. Pernambuco. Municipio de Camaragibe. Location: Aldeia.
HOLOTYPE (84-8401) and isotype (84-8402 & 84-8403) specimens deposited at URM
(Recife, Brazil); isolated from field samples (10 June 1997) and established in a pure
culture maintained at URM and INVAM; isotypes 84-8404& 84-8405 deposited at
Z+ZT (accession number ZT Myc 642), and 84-8406 & 84-8407 deposited at OSC (OSC
#134’504).
ETyMoLocy: heterogama referring to Endogone heterogama with which this new species
has been very often confused.
COMMENTARY: A detailed English species description of FE heterogama is not
needed, since several isolates of the new species were intensively studied during
the last 20 years (e.g. Franke & Morton 1994, Jeffries et al. 2007). Spores are
globose (165-280 um in diam) to subglobose to oval (159-265 x 195-295 um)
to rarely irregular, reddish-brown to dark red-brown to dark brown, with three
spore walls (ow, Mw and Iw). The ow three-layered (OWL1-3: OWL1 hyaline,
semi-persistent to persistent, 1.1-2.1 um thick, with papillae projections, 0.9-
1.5(—2.5) um in diameter and at 0.8—2.4 um distance from each other; ow12 red- |
brown to dark brown, laminate. 5.5-8.5 um thick, owL3 subtile, concolorous with
OWL2; Mw hyaline, bi-layered, in total 2.1-2.8 um thick; rw bi-to-tree-layered,
in total 2.1-3.4 um thick; owL2 & OWL3 and IWL2 staining dark red-brown and
purple, respectively, in Melzer’s reagent. Sporogenous cell globose to oblong,
32-51 x 31-45 um; germ shield yellow-brown to brown, simple, bi-lobed, oval
to ovoid to violin-shaped (75-105 x 55-75 um) to rarely irregular, generally
with 2 lobes rarely showing 4 folds; shield periphery generally smooth, rarely
with scarse dentate structures. Forms typical arbuscular mycorrhiza without
intraradical vesicles but extraradical auxiliary cells in the hyphal mycelium.
Fuscutata heterogama can be easily distinguished from D. heterogama by spore
color and especially by the morphology of the shield (see above; compare FIGS.
45-46 with Fic. 44). It can be easily distinguished from other Fuscutata spp. by
the ornamentation on the outer spore surface (see identification key below).
DISTRIBUTION: Fuscutata heterogama has a major distribution in warmer
climates (Mediterranean, Subropical and Tropical climates) but was confused
with Endogone heterogama which might have a less broad distribution. The
new species is frequent in Southern America as we have found it several times
in Colombia and Brazil. It was recently found also in Benin (West Africa;
Hountondji & Oehl, unpublished, Tchabi et al. 2008).
OTHER SPECIMENS EXAMINED: COLOMBIA. Eastern Plains. Sieverding collection.
UNITED STATES. Florida. Illionois. Specimens deposited at INVAM, and specimens
prepared by E Oehl during his visit at INVAM in 2002 (slides deposited at Z+ZT).
Specimens of the Embrapa Agrobiologia germplasm collection, Seropédica, Brazil,
New taxa in the Glomeromycetes ... 347
CNPAB2. BENIN: Guinea and Sudan savannas under yam and peanuts (Tchabi et al.
2008).
Fuscutata rubra (Stiirmer & J.B. Morton) Oehl, EA. Souza & Sieverd.,
comb. nov. (FIG. 47)
MB 511965
BasionyM: Scutellospora rubra Stirmer & J.B. Morton, Mycol. Res. 103: 951. 1999.
Fuscutata savannicola (R.A. Herrera &Ferrer) Oehl, EA. Souza & Sieverd.,
comb. nov. (FIG. 48)
MB 511966
BASIONYM: Gigaspora savannicola R.A. Herrera & Ferrer, Rev. Jard. Bot. Nacional,
Habananies /a19381-
= Scutellospora savannicola (R.A. Herrera & Ferrer) C. Walker & EE. Sanders,
Mycotaxon 27: 180. 1986.
Fuscutata trirubiginopa (X.L. Pan & G.Yun Zhang) Oehl, F.A. Souza & Sieverd.,
comb. nov.
MB 511967
BastonyM: Scutellospora trirubiginopa X.L. Pan & G.Yun Zhang, Mycosystema 16:
169.
Quatunica F.A. Souza, Sieverd. & Oehl, gen. nov.
MB 511976
Sporae terminaliter efformatae anguste adiacetae ad cellulas sporogeneas, quartuoribus
tunicis; scutellum germinale coniunctum ad tunicam interiorem, flavo-brunneum ad
brunneum, cum 8-30 compartimentibus et depressionibus germinationis; formans
structuras mycorrhizarum arbuscularum.
ETyMo_oey: From the Latin: ‘qua abreviation from quartuor (four) and ‘tunica’ (wall)
referring to the four spore walls, a unique character in Glomeromycota.
KEY CHARACTERS: Sporocarps are unknown. Spores singly formed on bulbous
sporogenous cells that are formed terminally on a subtending hypha that arises
from mycelia hyphae. Spores have 4 walls. Outer spore wall three- to four-
layered and continuous at least in part with the wall of the sporogenous cell;
pore between the spore and sporogenous cell is narrow and usually closed by
a plug formed by outer spore wall material. Two hyaline middle walls and a
hyaline inner wall form de novo during spore formation. Germination shield
generally arising on the outer surface of the inner wall or beneath a thin outer
layer of the inner wall, yellow-brown to brown, generally ellipsoid to oval,
or rarely reniforme to cardioforme, usually with a many (8-30) large folds
separating the shield into 8-30 ‘small compartments’; each small compartment
generally with one germ tube initiation (‘gti). From one or few gti germ tubes
may arise penetrating the middle walls and the outer wall; the periphery of
shield wall usually conspicuously dentate in planar view. One to several septa
often formed in the subtending hypha in some distance to the sporogenous cell.
348 ...Oehl, de Souza & Sieverding
Auxiliary cells formed in hyphal mycelium as far as they are known knobby
without spines on the surface. Forming typical arbuscular mycorrhizae, vesicle
formation in roots so far unknown.
TYPE SPECIES: Quatunica erythropus (Koske & C. Walker) F.A. Souza et al.
Key to Quatunica species
1a Spores white-yellow, to yellow-brown, 340-640 ttm in diam
TE eT oie Q. scutata (S. scutata sensu Silva et al. 2006b) [non editum|]
1b Spores orange-brown to dark red brown, 200-650 um in diam....... Q. erythropus
Quatunica erythropus (Koske & C. Walker) F.A. Souza, Sieverd. & Oehl,
comb. nov. (PLATE 9)
MB 511977
BASIONYM: Gigaspora erythropus Koske & C. Walker, Mycologia 76: 250. 1984.
= Scutellospora erythropus (Koske & C. Walker) C. Walker & EE. Sanders,
Mycotaxon 27: 181. 1986.
Discussion
Molecular analyses, based almost exclusively on 18S rRNA genes, have provided
the means to describe the phylum Glomeromycota and to reorganize almost
entirely the glomeromycotan phylogeny (Schtfsler et al. 2001). This resulted in
a new order, the Diversisporales, validly described by Walker & Schiifsler (2004),
which contains species and genera that can be distinguished morphologically
by the different spore formation types: acaulosporoid, entrophosporoid,
gigasporoid or glomoid. Such spore formation differences earlier served as
the basis for higher order ranks (Morton & Benny 1990). Only recently, new
families and genera have been proposed for the Glomeromycetes based on
combined molecular genetic and spore morphological data. Oehl & Sieverding
(2004) separated the Pacispora genus from Glomus in the Glomeraceae, order
Glomerales, based on morphological and germination characteristics, and
Walker et al. (2004) cited molecular data in describing the new Pacisporaceae
family in the order Diversisporales (Walker & Schiiler 2004). Sieverding & Oehl
(2006), Spain et al. (2006) and Walker (2007) also referred to molecular data to
review Entrophospora and the Archaeosporaceae, which prompted a taxonomic
reorganization of new fungal genera and families in the Archaeosporales based
on 18S and ITS rRNA-generated phylogenies (Walker et al. 2007).
We used published DNA sequences available from 18S and 25S rDNA for a
total of 17 former Scutellospora spp. and 7 Gigaspora spp. (FIGs. 10-11) and
morphological data from all previously described Scutellospora species as
a basis for separating the new families and genera in the Diversisporales
from the Gigasporaceae. The molecular analyses showed that Gigasporaceae
349
New taxa in the Glomeromycetes ...
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are phylogenetically closer to Racocetraceae and Dentiscutataceae than to
Scutellosporaceae and suggested that Scutellosporaceae is basal among the
four families that produce spores from a bulbous sporogenous cell. Contrary
to the evolutionary hypotheses expressed by Morton (1990) and Morton &
Benny (1990), Gigasporaceae may well have co-evolved with Racocetraceae
and Dentiscutataceae. Morphologically, however, Gigasporaceae differ as
greatly from Racocetraceae and Dentiscutataceae as Pacisporaceae differs from
Glomeraceae.
In this study, germination structures and spore wall composition were the
morphological characters that were supported by the molecular data. On
their basis the species could be organized within new or re-described genera
and families. This was easily done for Gigaspora, the only genus in the
Gigasporaceae, as the Gigaspora spore has only one wall and no germination
shield. However, it was more difficult to classify species forming a germination
shield. While striking genetical differences between the Scutellospora species
have become recognized (e.g. Simon et al. 1992, Helgason et al. 1998, Lanfranco |
et al. 1999, de Souza et al. 2005), researchers apparently have encountered
difficulties in grouping the species along genetic lineages. A recently
published key (Silva et al. 2006b) still focuses exclusively on spore size, spore
wall ornamentation, and wall characteristics. For the most part, the known
morphological differences in germ shields (Walker & Sanders 1986, Spain et
al. 1989) have been ignored in species taxonomies, so that this represents the
first time that germ shields have been observed, photographed, and used for
taxonomic classification. Little is known about the ontogeny of germination
shields of the different genera, which has been actually only described for
S. calospora by Walker & Sanders (1986). We do not know about the ontogeny
of shield formation of Racocetra, Cetraspora, Fuscutata and Dentiscutata, nor
is this knowledge necessary to identify the genera. Germ shield morphology
alone is sufficient clear, and stable.We confirmed that germ shield morphology
does not change with spore age for all examined species, as we could examine
numerous spores from different populations and sometimes very different
ecosystems (see TABLE 1: S. calospora, S. aurigloba, R. fulgida, R. gregaria,
R. verrucosa, C. pellucida, F. heterogama, D. reticulata, D. biornata).
Researchers who attempt to determine colors of very old field spores and shapes
when germination shield is hidden or incompletely visible may encounter
difficulties, particularly as the germ shield can readily be observed only when
seen in planar view. Spain et al. (1989) describe how to reveal germ shield
structure better by slightly moving the cover slide on a not yet completely
hardened fixative and by separating spore walls from crushed spores under the
stereomicroscope. There are a few species where the color of the germ shield
New taxa in the Glomeromycetes ... 355
may be ambiguous because parts of the originally hyaline shield brown slightly
over time on specific storage media (as in R. coralloidea and C. gilmorei) or
the spore wall turns so dark brown that the shield can no longer be seen (as in
S. savannicola herbarium specimens). Nonetheless, the germ shield folds are in
most cases easy to recognize, particularly when the shield is either very simple
(e.g. Scutellospora spp.) or very complex (Dentiscutata spp.). Some experience
is needed to identify the wavy folded, hyaline, multi-compartmented shields
of Racocetra and Cetraspora (FIG. 5 + PLATES 5-6). In such shields it is often
easier to count the number of germ tube initiations (gti) within the shield
compartments. Determining the number of middle walls may also confuse,
particularly during separation when the inner lamina of the outer wall can
be mistaken for a middle wall, making differentiation between Racocetra
and Cetraspora (or Dentiscutata and Quatunica) difficult. Here it is a matter
of crushing several spores so as to determine whether the wall layers stick
together or are easily separated. Finally, other spore features — size, color, wall
ornamentation, swelling of wall layers in PVLG based mountants, staining in
Melzer'’s reagent — may also be used to identify the species (and thus genus)
morphologically, as done previously (Silva et al. 2006b). Our own experience
with the new keys presented in this paper convinces us that identification of
species with bulbous sporogenous cells will be much easier now that germ
shields are used as diagnostic characters. Nonetheless, we suspect that the
genera presented here may contain additional groups with diverging shield
characters and possibly divergent phylogenies that may justify — in the future
— erection of new genera based on the number of lobes and gti or due to shield
composition (e.g. in Scutellospora and Dentiscutata, TABLE 4).
During our study we faced several difficulties related to the availability and
quality of live and herbarium specimens. Pure cultures were either unavailable
for most of the as-yet unsequenced species, (e.g. R. minuta, R. alborosea,
E savannicola) or pure cultures got lost with time (e.g. D. biornata). In a few
cases the herbarium type material was so poor in both spore number and spore
quality after 20-40 years of storage (e.g. for R. minuta, R. alborosea, and other
species described before 1990) that germ shields could not be found in the type
material. In a few other cases, the spores mounted or preserved in the type
material had obviously been immature, since the germ shield and sometimes
even the inner wall could not be detected on the majority of the spores. In
such cases we had to rely for the identification of shield morphology either on
the original species description or on our own specimens collected in different
parts of the world. Some species, e.g. R. alborosea and R. weresubiae, may be
conspecific but it was not our aim to correct this. We also noted that some
species should be redescribed (e.g. S. tricalypta), also not the goal of this paper.
There was, nonetheless, an urgent need to separate the species presented in the
356 ...Oehl, de Souza & Sieverding
type specimens for S. heterogama (Nicolson & Gerdemann 1968) from those
erroneously identified as such. Germ shield morphology clearly showed that
the two ‘heterogama’ species belong to two different genera, Fuscutata and
Dentiscutata (see description above).
In conclusion, the previously most commonly used morphological spore
characters like spore size, color and staining of different spore walls in Melzer’s
reagent or expanding of wall layers in lactic acid based mountants (Franke &
Morton 1994, Morton 1995) could not explain the rDNA-based phylogenetic
clades (PLATES 2-3; also Walker et al. 2004, de Souza et al. 2005, Ahulu
et al. 2006, Redecker et al. 2007). We have not investigated for this study
whether there are differences in the staining behaviour of the mycelium in
the roots among our new segregate genera. Such differences are important in
Paraglomeraceae and Archaeosporaceae (Morton & Redecker 2001), but we
found no literature indicating that this is the case in Gigasporaceae. We also did
not investigate whether there are specific differences between the genera in the
morphology of their auxiliary cells in the external root mycelium. Walker &
Sanders (1986) noted that such differences are only minor in Scutellospora and
then only consistent in differentiating Gigaspora from Scutellospora. Also, color
of the mycelium in soil may be different: some species (e.g., E heterogama,
D. reticulata) form light brown to brown walled mycelium while others, like
R. castanea, form mycelia with hyaline walls (pers. obs.). Information on all these
characters is available for only a few species and was considered unnecessary
to reorganize the genera presented here. Nevertheless, we propose that living
pure cultures be established and that detailed studies of the auxiliary cells
on extraradical mycelia and the ontogeny of germination shields in different
genera be carried out; studies of genetical and ecological characters are also
recommended. Unfortunately, much basic biology remains unknown in the
Glomeromycetes, including germination, the process so intimately linked with
the fitness and survival of these obligatory biotrophic symbiotic fungi. Although
axenic cultures may prove useful in the future, we believe that the important
spore characters must be observed in soil-grown spores that represent spores
found under field conditions that ecologists and taxonomists encounter in real
life.
Acknowledgments
We especially thank Joseph B. Morton and William Wheeler (West Virginia State
University; Morgantown, USA) for giving F. Oehl the opportunity to investigate, during
his visit in 2002, many Scutellospora and Gigaspora isolates maintained at INVAM. We
also acknowledge Joey Spartafora and Richard Halse for giving the possibility to visit
twice the OSC herbarium in Corvallis (Oregon) in 2002 and 2005. We are also thankful
to OSC (Oregon State University) and to FH (Harvard University) for providing us
SS
New taxa in the Glomeromycetes ... 357
with type and non type material of several Scutellospora species. James M. Trappe
(USDA-Corvallis, Oregon, USA), Janusz Blaszkowski (Department of Plant Pathology,
Adademy of Agriculture, Szeczin, Poland), José Oswaldo Siqueira (UFLA, Brazil), Gisela
Cuenca (IVIC, Venezuela), Leonor C. Maia, Bruno T. Goto & Thiago Morais (UFPE,
Brazil), Claudia Castillo (Universidad Catdélica de Temuco, Chile), Samar Velho da
Silveira (Departamento de Horticultura e Sivicultura, VERGS, Brazil) and Luis Rodrigo
Pérez-Camacho (AGRUCO, Universidad Mayor de San Simon, Cochabamba, Bolivia)
provided type or non type specimen of several Scutellospora species. Fabien Hountondji,
Atti Tchabi and Louis Lawouin (International Institute of Tropical Agriculture, Benin)
provided soil samples and specimen from their country. Kurt Ineichen is cordially
acknowledged for designing the illustrations in TABLE 2. This study was in part supported
by the Swiss Agency for Development and Cooperation (SDC) in the frame of the
Indo-Swiss Collaboration in Biotechnology (ISCB) programme, by the Swiss National
Science Foundation within the Programme NFP48 ‘Landscapes and habitats of the Alps;
by the Swiss Center for International Agriculture (ZIL) within the RFPP programme,
by Inter American Institute for Global Change Research grant IAI CRN II/14 which
is supported by the US National Science Foundation (Grant GEO-04523250) and for
the Brazilian National Council for Scientific and Technological Development (CNPq)
grant 477094/2004-0). We are grateful for Dra Gisela Cuenca (IVIC, Centro de Ecologia,
Caracas, Venezuela) and Dr. Gladstone Alves da Silva (Universidade Federal de
Pernambuco, Recife, Brazil) for reviewing the manuscript and making helpful comments
and suggestions. We appreciate the corrections by Shaun Pennycook, Nomenclatural
Editor, and suggestions by Lorelei L. Norvell, Editor-in-Chief.
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398-400.
Walker C, Sanders FE. 1986. Taxonomic concepts in the Endogonaceae: II. The separation of
Scutellospora gen. nov. from Gigaspora Gerd. & Trappe. Mycotaxon 27: 169-182.
Walker C, SchiiSler A. 2004. Nomenclatural clarifications and new taxa in Glomeromycota. Myc.
Res. 108: 981-982.
Walker C, Blaszkowski J, Schwarzott D, SchiiSler A. 2004. Gerdemannia gen. nov., a genus separated
from Glomus, and Gerdemanniaceae fam. nov., a new family in the Glomeromycota. Mycol. Res.
108: 707-718.
Walker C, Cuenca G, Sanchez F. 1998. Scutellospora spinosissima sp. nov. a newly described
Glomalean fungus from acidic, low nutrient plant communities in Venezuela. Annals Bot. 82:
721-725.
Walker C, Gianinazzi-Pearson V, Marion-Espinasse H. 1993. Scutellospora castanea, a newly
described arbuscular mycorrhizal fungus. Cryptogamie 14: 279-286.
Walker C, Vestberg M, Demircik FE, Stockinger H, Saito M, Sawaki H, Nishmura I, Schiissler A. 2007.
Molecular phylogeny and new taxa in the Archaeosporales (Glomeromycota): Ambispora fennica
gen. sp. nov., Ambisporaceae fam. noy., and emendation of Archaeospora and Archaeosporaceae.
Mycological Research 111: 137-153.
MY COTA YON
Volume 106, pp. 361-370 October-December 2008
Scutellospora pernambucana, a new fungal species in the
Glomeromycetes with a diagnostic germination orb
DANIELLE KARLA ALVES DA SILVA’, NICACIO DE OLIVEIRA FREITAS!
GISELA CUENCA’, LEONOR CosTA MAIA! & FRITZ OEHL?
'*leonorcmaia@yahoo.com.br
‘Departamento de Micologia, CCB, Universidade Federal de Pernambuco
Av. Prof. Nelson Chaves s/n, Cidade Universitaria, 50670-420, Recife, PE, Brazil
“Instituto Venezuelano de Investigaciones Cientificas, IVIC, Centro de Ecologia
Apdo. 21827, Caracas 1020 A, Venezuela
°Zurich-Basel Plant Science Center, Institute of Botany, University of Basel
Hebelstrasse 1, 4056 Basel, Switzerland
Abstract — A new species of the arbuscular mycorrhiza-forming Scutellosporaceae
(Glomeromycetes), described here as Scutellospora pernambucana, forms globose
to subglobose (rarely oval), yellow to brown-yellow glomerospores terminally on
sporogenous cells. The 110-150 um diameter spores have three unornamented spore
walls: outer (3-layered), middle (2-layered), and inner (3-layered). The germination
shield formed on the outer surface of the inner wall is circular, hyaline to (rarely) light
yellow, and has a characteristic coiled, orb-like structure formed by a single lobe that
surrounds the initial germ hole and terminally bears one germ tube initiation. This unique
germ shield structure easily distinguishes the new species from all other Scutellospora
species; S. projecturata, which forms a similarly coiled germ shield, is differentiated by
conspicuous columnar projections on the outer spore wall. S. pernambucana, collected
from the “Mata Atlantica’ and Caatinga biomes in Pernambuco State and a reforestation
area in Maranhao State, appears to be a common fungus in Northeastern Brazil.
Key words — Gigasporaceae, Glomeromycota, Diversisporales
Introduction
Arbuscular mycorrhizal fungal species that form spores on bulbous sporogenous
cells have been reorganized into four families based on germination structure
characteristics and phylogenies generated by independent analyses of 18S
and 25S rRNA gene sequences (Oehl et al. 2008). Species with only one spore
wall containing a warty inner germinal wall layer remain in the revised family
Gigasporaceae (Morton & Benny 1990, Oehl et al. 2008), while those producing
spores with 2-4 walls and a discrete germination shield on the innermost wall
362 ... Silva & al.
were transferred into three new families: a) species with a yellow-brown to |
brown germ shield were transferred into the Dentiscutataceae, b) species with |
a multiple-lobed, hyaline to subhyaline germ shield and multiple germ tube |
initiations were attributed to the Racocetraceae, and c) species with a simple, bi-
lobed to (rarely) mono-lobed, hyaline to subhyaline germ shield with (1-)2 germ ©
tube initiations were transferred to the Scutellosporaceae (Oehl et al. 2008). The —
newly established Scutellosporaceae is mono-generic with Scutellospora (Walker |
& Sanders 1986) as type genus. :
A new arbuscular mycorrhizal fungal species was found during diversity |
studies in Northeastern Brazil in Pernambuco State in a Tropical Atlantic |
forest fragment and in the Caatinga biome and in Maranhao State in a forest |
revegetation area. The species that forms spores terminally on sporogenous cells |
and a unique germination shield on the outer surface of the innermost spore |
wall is hereafter described under the epithet Scutellospora pernambucana.
i
}
|
|
|
Material and methods © /
Soil sampling and culturing of AM fungi
Soils were sampled in a tropical forest fragment (i.e., “Zambana”) situated |
among other native forest fragments and sugarcane plantations near the |
community of Igarassu, Pernambuco State, Northeastern Brazil. The forest |
site is located at 07°41°29"-07°43°17°S and 35°00°09"-34°58°38" W (20-110 m |
above sea level). In December 2007 samples were taken from the rhizospheres |
(0-20 cm depth) of several native plant species characteristic of the ‘Mata’
Atlantica’ tropical forest [e.g., Coccoloba latifolia (Polygonaceae), Maytenus
spp. (Celastraceae), Clusia nemorosa (Clusiaceae), Margaritaria nobilis |
(Euphorbiaceae), Guapira spp. (Nyctaginaceae), Calyptranthes brasiliensis
(Myrtaceae), Alseis pickelii (Rubiaceae), Pouteria peduncularis (Sapotaceae)].
The pH (H,O) of the sandy soil (91% sand) was 4.8, and available P (extracted | |
after Mehlich; Nelson et al. 1953) was low (3 mg kg").
The spore material was studied from field samples that were also used for pot | |
cultures with Panicum miliaceum as host. The pots (300 g) were filled with each —
soil sample and maintained at the greenhouse of the Department of Mycology, |
Universidade Federal de Pernambuco, Recife. No host colonization or spore |
formation has yet been observed.
Morphological analyses
Glomerospores were extracted from field soils by wet sieving (Gerdemarill
& Nicolson 1963) and sucrose centrifugation (Jenkins 1964). The spores were |
thereafter mounted in PVLG, PVLG + Melzer’s reagent, and in water, respectively .
(Brundrett et al. 1994). For the species descriptions, the terminology recently |
applied for other genera of the Diversisporales (Oehl & Sieverding 2004, Oehl et”
Scutellospora pernambucana sp. nov. (Brazil) ... 363
al. 2006, Sieverding & Oehl 2006, Palenzuela et al. 2008) were adopted, as well
as the terminology of Walker & Sanders (1986) for the germ shield structures.
For the spore denomination the terminology proposed by Goto & Maia (2006)
was used.
Description of the new species
Scutellospora pernambucana Oehl, D.K Silva, N. Freitas, L.C. Maia, sp. nov.
MycoBank MB 512130 FIGS. 1-10
Sporocarpia ignota. Sporae singillatim in solo efformatae anguste adiacetae ad cellulas
sporogeneas subterminales vel intercalares, flavae ad bruneo-flavae ad flavo-bruneae,
globosae (110-145 um in diametro) vel subglobosae vel ovales (105-135 x 120-150 um);
sporae tunicis tribus: tunica exterior stratis tribus, in totum 3.5-5.5 um crassa in aqua,
expandens cum pressione (5.5-8.5 um crassa) vel in PVLG (6.0-)9-14 um crassa; stratum
exterius tunicae exterioris hyalinum, evanescens ad semi-persistens, 1.0-2.lum crassum;
stratum medium laminatum, flavum ad brunneo-flavum, expandens cum pressione in
aqua vel in PVLG; stratum interius tunicae exterioris flavum ad brunneo-flavum, subtile,
0.5-1.1 um crassum; tunica media et tunica interior de novo formantes stratis hyalinibus;
tunica media duobus stratis in totum 1.1-2.6(-3.3) um crassa; tunica interior tribus
stratis, 1.7-2.8(-3.5) um crassa; solo stratum medium tunicae interioris purpureo colorans
reagente Melzeri; scutellum germinale coniunctum ad tunicam interiorem, hyalinum
vel rarum albo-flavum; circulare vel ovale, 50-80(-90) x 60-80(-94) um in diametro,
cum lobo singulare, orbem formans, generaliter cum depressione singulare germinationis;
structurae mycorrhizarum ignotae.
HOLOTYPE: 83-8301 (URM No. 79239) from ‘Zambana tropical forest fragment, Usina
Sao José, Igarassu, Pernambuco State, Brazil.
ETYMOLOGY: pernambucana referring to the State in Northeastern Brazil where the new
species was found first in two different biomes.
SPOROCARP FORMATION — unknown.
GLOMEROSPORES are singly formed in soils terminally on a subterminal or
intercalary bulbous suspensor cell (= ‘sporogenous cell; Fic. 1). Glomerospores
are dark yellow to brown yellow to yellow brown, globose (110-145 um in
diameter) to subglobose (105-135 x 120-150 um) to rarely oval, and have with
three walls: an outer, a middle and an inner wall (ow, Mw and Iw; FIG. 2).
OUTER WALL is in total 3.6-5.5 um thick in water, but expanding under
pressure to 5.5-8.5 um and to (6.0—-)9.0-14 um in PVLG based mountants;
Ow consists of three layers (FIGS. 3-7): outermost wall layer (OWL1), hyaline to
subhyaline to light yellow, evanescent to semi-persistent and about 1.0-2.1 um
thick (FIGs. 3, 4, 7). Under field conditions, owL1 is sometimes recognized only
as fragmented remnants on the spore surface (FIG. 6). The second layer (owL2)
is dark yellow to brown-yellow, 2.0-3.6 um thick in water, expanding to 5.0-7.5
lum under pressure and to (7.0—-)8.5-12.0 um in PVLG based mountants (FIGs.
1-6); owL3 is concolorous with owr2, or slightly lighter in color, thin and
flexible (0.5-1.1 um thick) and usually difficult to observe as closely adherent
364 ... Silva & al. |
|
to laminate OwL2; when separating from OWL2, OWL3 often shows several folds.
(FIG. 5). None of the layers stain in Melzer’s reagent. The straight pore channel
at the spore base (about 2.8—5.2 um broad) is often closed by a plug formed by
spore wall material of owL2, but sometimes appears to be open. )
MIDDLE WALL consists of two hyaline, flexible to semi-flexible layers (IGS.
4-6). Outer layer (MwL1) is about 0.6-1.3 um thick. Inner layer (Mw12). |
generally is more rigid as slightly thicker, 1.0-1.5(-2.0) um. Both layers mal
slightly expand in PVLG based mountants.
INNER WALL is three-layered (FIG. 4-6) bearing a germination shield on the |
outer surface (FIGS. 6-7). Outer layer of the inner wall (Iw11) is hyaline, semi-
flexible to sometimes finely laminate and 0.4-1.0 um thick. Second layer (Iw12)
is unite to finely laminate, 1.5-2.0(-2.7) um thick. Innermost layer (IWL3) is |
thin (0.4-0.8 tm thick), flexible, and — as usually tightly adherent to Iw12_
— often difficult to observe. The three layers may slightly expand in PVLG_
based mountants. Only Iw12 stains in Melzer’s reagent become dark red purple: |
within a few hours.
SPOROGENOUS CELL is globose to elongate, concolorous with the spore, or
slightly lighter or darker in color, and 31-48 um long and 26-43 um broad |
(FIGS. 1-3). The two wall layers generally visible on the sporogenous cell are |
continuous with owL1 and OWL2. OWL1 on the sporogenous cell is about.
0.7-1.4 tm and evanescent to semi-persistent; persistent OWL2 is about)
2.0-3.5 um thick. The sporogenous hypha attached is 8-11 um broad and also |
bi-layered tapering to 5-7 um within 200-350(-500) um distance from the
sporogenous cell. The sporogenous hyphal wall is concolorous with the spore.
wall, or sometimes darker yellow than the spore, and tapers from 1.7-2.8 um to!
0.9-1.5 um within this distance. Several (3-7) septa originating from the inner |
|
layer (OWL2) might be visible in the hypha. |
|
GERMINATION SHIELD is hyaline to subhyaline to rarely light yellow, circular
(50—)62-85(-95) um to subglobose to rarely oval (50-75(-85) x 63-90(-98) |
uum). Shield with one initial germ hole (gh) in the shield center (FIG. 6-7; and.
drawings of FIGs. 11-14), and one lobe which surrounds one to two folds the.
shield center and forms a circular, orb-like shield structure (FIGs. 6-14). One .
germ tube initiation (gti, 2.2-4.0 um in diameter) is positioned towards the end
of the lobe, from where one to rarely two dark yellow germination tubes emerge’
during germination and penetrate the middle and the outer wall, and branch in’
a short distance from the spore (FIG. 9). Shield walls (0.8-1.5 um thick) often +
appear sparsely dentate and generally wrinkling (FIGs. 7-10), since the shield
wall respective the lobe wall is rather thin, 0.6-1.2 um.
SPORE DEVELOPMENT — The major spore developmental stages were deduced’
from clearly identified spores of S. pernambucana found in the field samples.
Scutellospora pernambucana sp. nov. (Brazil) ... 365
Figs. 1-10: Scutellospora pernambucana Figs. 1-3. Spores with three spore walls (outer, middle and
inner wall; OW, MW, IW), and formed terminally on sporogenous cells (sc). Sporogenous hyphae
with several initial, but rudimentary branchings (pegs), and with several septa. 2-3. Germination
shield visible in cross view on the inner wall. Figs. 3-7. Spore wall structure with three layered
OW (OWL1-3), bi-layered MW (MWLI1-2) and three-layered IW (IWL1-3). Fig. 6. IWL2 with
purple reaction in PVLG + Melzer’s reagent. Figs. 6-10. Orb-like germination shield formed on the
outer surface of IW. The initial germ hole (gh; Fig. 6-7) and the terminal germ tube initiation (gti;
Fig. 9) are not easily detectable. Fig. 8. Lobe apparently changed growth direction during shield
formation, but remained tightly attached to the shield. Fig. 9. Germ tube (gt) emerging from the
gti. Fig. 10. Germination shield isolated from IW.
366 ... Silva & al.
50 um
Figs. 11-14. Scutellospora pernambucana: drawings of circular to oval germination shields with
a central initial germ holes (gh), single lobes that terminally or subterminally bear one germ tube
initiation (gti) from where the germ tubes emerge during germination. On a few spores, the lobes
apparently changed the growth direction during shield formation, but remained tightly attached to
the shield (Fig. 11). The gti might not be detectable in young, developing spores (Fig. 12).
First the outer spore wall differentiates into one evanescent to semi-persistent
outer layer (OWL1), a laminate layer (OWL2), and the adherent thin inner layer
(Ow13). The bi-layered middle wall (mw) and (subsequently) the three-layered
inner wall (tw) develop de novo without visible connection with the outer wall.
Finally, the orb-like germination shield develops on the outer surface of the
inner wall.
AUXILIARY CELLS — unknown.
MycoORRHIZA FORMATION — unknown.
DISTRIBUTION — Brazil: so far found in the semi-humid Atlantic Forest or
‘Mata Atlantica biome, in the Municipality of Igarassu and in the Caatinga
biome, in the Municipality of Araripina (both in Pernambuco State) and in a
Forest regeneration area in Sao Luis (Maranhao State), suggesting that the new
fungus might be common in Northeastern Brazil.
OTHER SPECIMENS OR ISOLATES EXAMINED: BRAZIL. Pernambuco State. Igarassu,
Usina Sao José, Zambana fragment — 83-8302 (URM79240) & 83-8303 (URM79241)
Scutellospora pernambucana sp. nov. (Brazil) ... 367
deposited in Recife, Brazil; 83-8304 & 83-8305 deposited at OSC (No. OSC# 134503;
Corvallis, Oregon, USA); 83-8306 & 83-8307 deposited at Z+ZT (No. ZT Myc 641;
Zurich, Switzerland); 83-8308, 83-8309 & 83-8310 (Oehl collection). Araripina
— specimens deposited at URM. Maranhao State. Sao Luis — specimens deposited
at URM.
Discussion
Scutellospora pernambucana can easily be distinguished from all other known
species in Scutellospora by the spore wall structure and in particular by the
unique structure and shape of the mono-lobed germination shield resembling a
germination orb. Of all known Scutellospora species, only S. projecturata forms
a similar, mono-lobed, coiled germ shield, but the later species has distinctive,
prominent columnar protuberances on the spore surface (Kramadibrata et al.
2000) while S. pernambucana has no ornamentation on any of the spore walls.
Moreover, the S. projecturata shield has a smooth but firm shield wall (FIc. 12
in Kramadibrata et al. 2000), while the thin S. pernambucana shield wall is
conspicuously wrinkled. The pronounced expansion of the laminate owL2 in
lactic acid-based mountants or under pressure and the evanescent to semi-
persistent nature of OWL1, an extremely rare unique feature for Scutellospora,
are two further helpful diagnostic characters. Observation of these last features
is particularly helpful when the shield cannot be observed in planar view or has
not yet differentiated in young, developing spores. However, it is also possible
that S. projecturata has an evanescent outermost wall layer on the surface of
the laminated, structural wall layer that was not reported in the protologue. In
contrast to the two aforementioned species, most other Scutellospora species
(e.g. S. calospora and S. pellucida) have a unit, semi-persistent to persistent layer
on the outer surface of the structural, laminate layer (here owL2), (Koske &
Walker 1986).
We interpret the S. pernambucana germination shield type as simple-
structured and mono-lobed, since obviously only one lobe develops during
shield formation. The single lobe surrounds the germ shield initiation, i.e. the
initial germ hole (gh; Walker & Sanders 1986) in the shield center. In all shields
observed, the lobe surrounded the shield center one to two times and was thus
responsible for the characteristic orb-like shield structure (FIGs. 6-14). In a few
specimens it appeared that the lobe started to cover the shield center in one
direction but obviously changed the direction and thus did not form a typical
orb (FIGs. 8, 13). Kramadibrata et al. (2000) described the S. projecturata germ
shield as coiled but did not refer to the shield structure as an ‘orb’ or ‘orb-like’ In
that paper, the illustrated shield of a crushed spore somehow resembles the S.
pernambucana shield structure presented here in FIG. 13 for, so we assume that
the shield structures of both species are indeed closely related.
368 ... Silva & al.
There are several other Scutellospora species with simple and hyaline to
subhyaline germ-shields that are, however, bi-lobed (Silva et al. 2006b, Oehl
et al. 2008): e.g. S. calospora (Koske & Walker 1986), S. tricalypta (Ferrer &
Herrera 1981), S. dipapillosa (Koske & Walker 1985), S. arenicola (Koske &
Halvorson 1990), and S. aurigloba (Walker & Hall 1991). In spores of these
species, fully developed shields are violin-shaped, oval to ovoid, or (rarely)
cardioid, each generally with one germ tube initiation (gti) on the terminal
ends of the two lobes (see Walker & Sanders 1986). Other species such as
S. castanea (Walker et al. 1993), S. heterogama sensu Franke & Morton (1994)
and Jeffries et al. (2007), and S. reticulata (de Souza et al. 2005) have more
complex, multiple-lobed shields with multiple gti or yellow-brown to brown
shields that species-specifically may have multiple small compartments and
multiple gti, and a conspicuously dentate shield periphery. Such species have
been transferred - due to their shield characteristics and for phylogenetic
reasons — to newly described genera in the Racocetraceae and Dentiscutaceae
(Oehl et al. 2008). |
So far, germination orbs have been described for only a few Acaulospora and _
Kuklospora species of the Acaulosporaceae (Spain 1992, Sieverding & Oehl 2006)
but not for Scutellospora. Oehl & Sieverding (2004) did not correctly interpret
the circular characteristics of germination orbs (Spain 1992) when they called
the Pacispora spore germination structures germination orbs. The germination
structures in Pacispora species generally form first during germination and not,
as in Scutellospora, during late spore formation, and thus should not be even
called shields. Their germ structures are not orb-like but multiple-lobed, and
the germ tubes emerge, as in acaulosporoid spores of Ambispora (Spain et al.
2006, Goto et al. 2008), from the germination center between the lobes (Oehl
unpublished). Interestingly, the germ tubes in the phylogenetically related
Scutellospora, Acaulospora and Kuklospora (e.g. Schiifler et al. 2001, Silva et al.
2006a, Palenzuela et al. 2008) emerge from the germ tube initiations positioned
at the termina! ends of the lobed germination shields (‘gti in Walker & Sanders
1986, ‘germ tube loci in Spain 1992), and their germination structures generally
persist once formed during spore formation. After analyzing recently published
phylogenetic trees for Glomeromycota species (e.g. Santos et al. 2006, Silva et al.
2006a, Palenzuela et al 2008), we concluded that formation of a persistent germ
structure (= germ shield) might be an evolutionary advance in Scutellospora
and Acaulospora over other genera (e.g. Ambispora and Pacispora) that also
form an inner, so-called germinal wall. We further assume that formation of a
typical germ orb in S. pernambucana like that found in Kuklospora colombiana
(Spain et al. 1992, Sieverding & Oehl 2006) and K. kentinensis (Oehl, pers.
obs.) might be ancestral in Scutellospora, as is inferred for K. colombiana in
the Acaulosporaceae (e.g. Santos et al. 2006, Palenzuela et al. 2008). This later
Scutellospora pernambucana sp. nov. (Brazil) ... 369
hypothesis can be tested for S. pernambucana only after DNA is extracted from
viable spores isolated from the field or — even better -successful pure cultures
of the new fungus.
Acknowledgements
The authors acknowledge Marta Cabello (Instituto Spegazzini, La Plata, Argentina)
and Mauritz Vestberg (MTT Agrifood Research, Vihtavuori, Finland) for reviewing the
manuscript and making helpful comments and suggestions. Thanks are also due to Prof.
Ana Carolina Lins e Silva, who provided information regarding the plant species in the
Forest area. This work was supported by the Conselho Nacional de Desenvolvimento
Cientifico e Tecnolégico (CNPq) and Coordenacao de Aperfeigoamento de Pessoal de
Nivel Superior (CAPES) that provided, respectively, a Master scholarship to Danielle
Alves da Silva, a PhD scholarship to Nicacio de Oliveira Freitas and a research
grant to Leonor C. Maia. This work was also supported by the Universidade Federal
de Pernambuco, which provided a grant to F. Oehl as ‘visiting professor. This is a
contribution to the project “Sustainability of remnants of the Atlantic rainforest in
Pernambuco and its implications for conservation local development’, a Brazilian-
German scientific cooperation within the program “Science and Technology for the
Atlantic Rainforest” funded by CNPq (590039/2006-7) and BMBF (01 LB 0203 Al),
permitted and logistically supported by Usina Sao José $.A/Grupo Cavalcanti Petribu.
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Baza (Granada, Spain). Mycologia 100: 296-305.
Santos JC, Finlay RD, Tehler A. 2006. Molecular analyses of arbuscular mycorrhizal fungi colonizing
a semi-natural grassland along a fertilization gradient. New Phytol.172-159-168.
Schiifler A, Schwarzott D, Walker C. 2001. A new fungal phylum, the Glomeromycota: phylogeny
and evolution. Mycol. Res. 105: 1413-1421.
Sieverding E, Oehl F 2006. Revision of Entrophospora, and description of Kuklospora and
Intraspora, two new genera in the arbuscular mycorrhizal Glomeromycetes. J. Appl. Bot. Food
Qual. — Angew. Bot. 80: 69-81.
Silva GA, Lumini E, Maia LC, Bonfante P, Bianciotto V. 2006a. Phylogenetic analysis of
Glomeromycota by partial LSU rDNA sequences. Mycorrhiza 16: 183-189.
Silva GA, Maia LC, Stiirmer SL. 2006b. A dichotomous key to Scutellospora species using
morphological characters. Mycotaxon 94: 293-301.
Spain JL. 1992. Patency of shields in water mounted spores of four species in Acaulosporaceae
(Glomales). Mycotaxon 43: 331-339.
Spain JL, Sieverding E, Oehl F. 2006. Appendicispora, a new genus in the arbuscular mycorrhizal-
forming Glomeromycetes, with a discussion of the genus Archaeospora. Mycotaxon 97:
163-182.
Walker C, Hall IR. 1991. Lectotypification of Scutellospora auriglobosa (Glomales). Mycol. Res. 95:
398-400.
Walker C, Sanders FE. 1986. Taxonomic concepts in the Endogonaceae: III. The separation of
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Volume 106, pp. 371-378 October-December 2008
A new species of Leucoagaricus (Agaricaceae)
from Colombia
ADRIANA ORTIz"?*, ANA ESPERANZA FRANCO-MOLANO!
& MAURICIO BACCI JR.?
* adortizr@unal.edu.co, afranco@quimbaya.udea.edu.co, mbacci@rc.unesp. br
' Laboratorio de Taxonomia y Ecologia de Hongos, Instituto de Biologia
Universidad de Antioquia A.A. 1226, Medellin, Colombia
? Facultad de Ciencias, Universidad Nacional de Colombia
Codigo postal CO05-07, Medellin, Colombia
° Molecular Evolution Laboratory, Center for the Study of Social Insects
Sao Paulo State University, Rio Claro, S.P, Brazil
Abstract — A new taxon of Leucoagaricus is proposed as L. amazonicus. The collection
representing this species was collected in the lowlands of the Amazonas Department,
Colombia, on a dump produced by ants belonging to Acromyrmex octospinosus. The
very particular habitat, the small size and dark brown basidiomata that stain yellowish
when bruised, the peronate inferior annulus, and the cheilocystidia which are relatively
short, capitate and filled with dense brown contents are some of the features of this
species.
Key words — Agaricales, taxonomy, Amazonia
Introduction
Leucoagaricus Locq. ex Singer belongs to the family Agaricaceae (Singer
1986, Vellinga 2004), and has approximately 75 species (Kirk et al. 2001). It
is characterized macroscopically by having small to medium size basidiomata
with lepiotoid or pluteoid habit that varies from fragile and slender to sturdy
and fleshy; a pileus surface that is radially fibrillose, floccose, squamulose to
fibrillose-scaly, and rarely granulose; an entire or very short striate margin; a thin
or thick context that may or may not stain when bruised; free, rarely collariate
lamellae; a central, equal to bulbous stipe, and a membranous, + persistent,
and sometimes moveable annulus. Micromorphologically, the Leucoagaricus
species have basidiospores with or (more commonly) without a rudimentary,
rarely distinctive, germ pore and a metachromatic endosporium; a rather thin,
smooth to ornamented spore wall; a regular to trabecular hymenophoral trama;
372 ... Ortiz, Franco-Molano & Bacci
a variable pileipellis that is either a trichoderm or a cutis of repent and radially
arranged hyphae lacking sphaerocytes; pleurocystidia that are sometimes
present; abundant cheilocystidia, and absence of clamp connections (Singer
1986, Vellinga 2001).
Leucoagaricus species are saprophytic, growing on soil, wood chips, and
sawdust in woods, grasslands, road verges, and dune vegetation. The genus
is widely distributed but represented by more species in the subtropical and
tropical areas than in cool-temperate regions, and it is absent from artic and
alpine areas (Vellinga 2001).
Previous molecular studies (Johnson & Vilgalys 1998, Johnson 1999, Moncalvo
et al. 2002, Vellinga 2004) imply that Leucoagaricus and Leucocoprinus Pat. form
a monophyletic clade. However, more systematic studies of tropical species
and species cultivated by attine ants are needed to resolve the phylogenetic
relationships of this group completely.
In Colombia the genus has been little studied, with only one species,
—————
Leucoagaricus rubrotinctus (Peck) Singer, a very common terrestrial fungus
formally reported (Franco et al. 2000).
Materials and methods
Specimens were collected, documented, photographed, and dried according to
Lodge et al. (2004). Color designations follow Kornerup & Wanscher (1983).
Line drawings were made with the aid ofa drawing tube. All micromorphological
structures from a tangential section were measured in 3% KOH. Abbreviations
describing the basidiospore size are n = number of spores measured, X = mean
length x mean width, Q = length/width ratio.
This study is based on 16 sequences. Genomic DNA was extracted from
the fresh Colombian material based on the methods of Silva-Pinhati et al.
(2004); other sequences used in the analyses were retrieved from GenBank.
The sequences were from: Leucoagaricus gongylophorus (Moller) Singer,
L. melanotrichus (Malencon & Bertault) Trimbach, Lepiota cf. abruptibulba (PA
156), Lepiotaceae sp. (PA 170), Lepiota cf. zamurensis (PA 415), Lepiotaceae
sp. (PA 280), Leucoagaricus meleagris (Sowerby) Singer, L. americanus (Peck)
Vellinga, L. bresadolae (Schulzer) Bon & Boiffard, and L. georginae (W.G. Sm.)
Candusso. Macrolepiota procera (Scop.) Singer was chosen as the outgroup.
GenBank access numbers are given in Fic. 3. The new sequence with the
accession number EU940371 was also deposited in GenBank. ;
Molecular studies were based on the ribosomal DNA region encompassing
the internal transcribed spacer 1 (ITS1), the 5.8S gene and the internal
transcribed spacer 2 (ITS2) regions. Angle amplicon containing these regions
was obtained through a polymerase chain reaction (PCR) with the primers ITS5
Leucoagaricus amazonicus sp. nov. (Colombia) ... 373
and ITS4 (White et al. 1990). Sequencing of strands was performed with an ABI
model 377 sequencer (Applied Biosystems, Foster City, CA) using a BigDye™
Terminator Cycle Sequencing Ready Reaction Kit (Applied Biosystems) and
with the same primers used for the PCR. The sequences were aligned with
Clustal W program (Thompson et al. 1994).
Maximum parsimony analysis of nrITS sequences were performed using
PAUP* version 4.0b10 (Swofford 2002). The final alignment had 774 characters
of which 231 were parsimonious informative.
One hundred heuristic searches were conducted with random sequence
addition and tree bisection-reconnection (TBR) branch-swapping algorithm,
collapsing zero-length branches and saving all minimal length trees (MulTrees);
gaps were treated as missing. To measure relative support for the resulting
clades, 1000 bootstrap replications (Felsenstein 1985) were performed under
the fast stepwise addition option. The parsimony analysis resulted in three
trees, with a tree length of 518 steps, with a consistency index (CI)= 0.7876 and
the retention index (RC) = 0.6683.
Taxonomy
Leucoagaricus amazonicus A. Ortiz & Franco-Mol., sp. nov. FiGuREs 1-2
MycoBAank MB 512219, GENBANK EU940371
Species nominate Leucoagaricus amazonicus descripta est per: singularitatem habitus
moris sui in sordio receptaculo speciei Acromyrmex octospinosus, per magnitudinem
parvam suam, per adustem colorem basidiocarporum pallorem contrahere quandocumque
manu operari est, etiam cheilocistidios perexiqui sunt, cum capitibus atque materie
contineretur obscurissumum colorem (similes fabarum coffearum color), et annulus ab
infimo peronatus est. Principia priora descripta species Leucoagaricus amazonicus ceteris
omnibus speciebus ab ipse genere discernerent.
EryMoLocy: Named for the Colombian Amazonia where this species was found.
Ho.otyPe: COLOMBIA. DPTO AMAZONAS: Municipio de Leticia. Parque Natural
Nacional Amacayacu, Isla de Mocagua, 3° 49’ 51.48” S; 70° 19° 59.58” W, flood plain
forest, December 11, 2003, A. Ortiz 58 (Holotype: HUA).
| Pireus 3-4 cm in diameter, convex to plano-convex; surface velvety when
_ young to velvety at the center and fibrillose-squamulose toward the margin with
age; center, fibrils and squamules dark brown (near 7F8), on a white surface in
| mature basidiocarps; margin incurved to decurved, eroded. PILEUS CONTEXT
3mm thick, white, changing to yellowish (444) when exposed. Odor and taste
not distinctive to slightly sweet. LAMELLAE up to 4 mm broad, white, changing
to yellowish (4A4) when bruising, free and remote from stipe, crowded; edge
| entire, marginate brown (7F8) near at the margin of pileus; lamellulae of different
lengths. Stipe 3 cm in length, up to 0.2 cm in diameter near the apex, central,
| slightly clavate; surface fibrillose-squamulose, concolorous with pileus, annulate.
_ STIPE CONTEXT white, stuffed. ANNULUS inferior, peronate, membranous, thin,
374 ... Ortiz, Franco-Molano & Bacci
Ficure 1. Basidiomata of Leucoagaricus amazonicus (Holotype). Photo: Ortiz A.
concolorous with stipe surface but darker at the external edge. SPORE PRINT not
obtained. Basip1ospores (Fig. 2D) 4.0-6.0(-7.0) x (3.0-)3.5-4.0(-5.0) um,
(n = 25; X = 5.12 x 3.98 um; Q = 1.41 + 0.03), ellipsoid, amygdaliform, hyaline |
in KOH, dextrinoid in Melzer’s reagent, metachromatic in cresyl blue after a |
treatment with ammonium and acetic acid, acyanophilic in cotton blue, spore
wall smooth and germ pore rudimentary to absent. Basip1A (Fic. 2C) 10-15 x |
4-6 um, subcylindric to narrowly clavate, with four sterigmata. LAMELLAE EDGE |
sterile. CHEILOCYSTIDIA (Fic. 2B) abundant, (15—)20-36(-43) x (8-)10-14
um, capitate, filled with dense brown content. PLEUROCYSTIDIA absent.
HYMENOPHORAL TRAMA regular, composed of parallel hyphae up to 8 um in|
diameter. SUBHYMENIUM pseudoparenchymatous and poorly differentiated.
PILEIPELLIS (Fic. 2A) a trichoderm of erect, densely crowded, unicellular hairs, |
up to 126 um long, and up to 17 um in diameter, and shorter elements, clavate, |
broadly clavate to pyriform, thin walled, hyaline, frequently filled with a dense
brown content. PILEAL TRAMA of interwoven, cylindrical to inflated hyphae,
3.2-12 um in diameter, thin walled, hyaline, often branched. STIPITIPELLIS
composed of vertically oriented and parallel cylindrical hyphae, up to 8 um in»
diameter, thin walled, hyaline, giving rise to abundant hairs similar to those of
the pileipellis. CLAMP CONNECTIONS absent. |
Leucoagaricus amazonicus sp. nov. (Colombia) ... 375
FiGure 2. Leucoagaricus amazonicus. Microscopic features taken from the holotype.
A. Pileipellis elements. B. Cheilocystidia. C. Basidia. D. Basidiospores. Scale bar = 10 um.
HABITAT AND DISTRIBUTION - Solitary to dispersed, growing on dumps
produced by ants belonging to Acromyrmex octospinosus (Reich 1793). These
ants build detritus piles away from their gardens and outside of their nest.
Leucoagaricus amazonicus has been observed only in this habitat.
Discussion
In its habitat, Leucoagaricus amazonicus resembles L. gongylophorus, from
which it differs in the absence of oleiferous hyphae, the presence of short,
capitate cheilocystidia filled with brown contents, bigger spores, and smaller
basidiomata. Furthermore, while L. gongylophorus is cultivated by leaf-cutting
ants belonging to Atta and Acromyrmex, L. amazonicus was found on a dump
produced by Acromyrmex octospinosus (Bononi et al. 1981, Muchovej et al.
1991).
_ Although L. amazonicus basidiospores have a rudimentary to absent germ
| pore, the white pileus context and white lamellae that yellow on exposure or
_ bruising and the pileus surface that breaks into fibrillose-squamulose scales
| place this species into section Annulati (Fr.) Singer, together with L. naucinus
376 ... Ortiz, Franco-Molano & Bacci
Macrofepiota procera
AF482848
Lepiota cf. abruptibulba (PA 156)
AF079732.4
68
Leucoagaricus amazonicus
400. Lepiotaceae sp. {PA 170)
AF079734
Leucocoprinus cf. zamurensis (PA 415)
7
62 AY176407
Lepiotaceae sp. (PA 280)
AFO79744
Leucoagaricus meleagris
400 AY 176407
Leucoagaricus méleagns
AF482867
Leucoagaricus bresadolae
AF295929
100
Leucoagaricus americanus
62 AF295929
Leucoagaricus americanus
98 AF295928
Leucoagaricus georginae
400 AY176413
Leucoagaricus georginae
AY 243634
Leucoagaricus mefanotrichus
AY176417
50
Leucoagaricus gongylophorus
AY642815.1
Leucoagaricus gongylophorus
AY642811,1
UG
Leucoagaricus gongylophorus
AY642816.1
FrGureE 3. One of the three most parsimonious trees based on nrITS data. Bootstrap values = 50%
are given in bold on the branches. GenBank accession numbers are given for each collection.
Leucoagaricus amazonicus sp. nov. (Colombia) ... 377
(Fr.) Singer, L. leucothites (Vittad.) Wasser, L. olgae (Velen.) M.M. Moser, and
L. gongylophorus (Singer 1986, Candusso & Lanzoni 1990). However, while
L. amazonicus grows on ant dumps, the other species in the section grow in
grassy areas on disturbed ground or, in the case of L. gongylophorus, on ant
nests. Microscopically, none of the species in the section combine the characters
of L. amazonicus, which includes the short and capitate cheilocystidia filled
with dense brown contents.
The combination of the colored cheilocystidia, rudimentary to absent germ
pore and the yellowing reaction suggest a resemblance to L. americanus or
similar species such as L. meleagris and Lepiota besseyi H.V. Sm. & N.S. Weber.
However, these species are characterized by having bigger cheilocystidia with a
pronounced mucronate apex, and bigger spores with a germ pore (Pegler 1983,
Smith & Weber 1987, Vellinga 2000). L. amazonicus morphologically resembles
some species belonging to section Piloselli (Kiuhner) Singer (Reid 1990), from
which it differs by tissues that do not become green in KOH.
Analyses of the ITS data set place L. amazonicus with a 100% bootstrap support in
a clade with Lepiota cf. abruptibulba (PA 156), Lepiotaceae sp. (PA 170), Lepiota
cf. zamurensis (PA 415), Lepiotaceae sp. (PA 280) according to the information
obtained from the GenBank (Fic. 3). However, morphological description of
these species is not available, limiting our comparison to sequence analyses. A
100% bootstrap supports the clade of Leucoagaricus amazonicus and its closest
relative, which is an undescribed species cited as Lepiotaceae sp. (PA 170) by
Mueller et al. (1998). Leucoagaricus amazonicus does not cluster with other
species with dark brown cheilocystidia, such as those in the L. americanus
group or with species (such as L. georginae) in section Piloselli.
Acknowledgments
This study was possible due to the support of CODI, Universidad de Antioquia, to Ana
Esperanza Franco M., FAPESP and CNPg to Mauricio Bacci Jr., and NWO and WOTRO
grant WB 84-525 to Dr. Teun Boekhout (CBS, Utrecht, The Netherlands). The authors
wish to acknowledge Carlos Alberto Lopez Quintero, Felipe Cardona, Aida Vasco,
Leonel Panduro and Jorge Navarro for the field assistance. Thanks are extended to Drs
Laura Guzman and Else Vellinga for the critical review of the manuscript and to Javier
Munoz for the Latin diagnosis.
Literature cited
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formigueiro de Atta sexdens rubropilosa Forel. Rickia 9: 93-97.
Candusso M, Lanzoni G. 1990. Lepiota s.l. Fungi Europaei 4. Libreria Editrice Giovanna Biella,
Saronno, Italy. 743 pp.
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y otros hongos). Colciencias, Universidad de Antioquia, Medellin, Colombia. p. 30.
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Kirk PM, Cannon PF, David JC, Stalpers JA. 2001. Dictionary of the fungi. 9° ed. CAB International,
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pp. 30 pl.
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leaf-cutting ants. Mycol. Res. 95 (11): 1308-1311.
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2034-2034.
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Reid DA. 1990. The Leucocoprinus badhami complex in Europe: species which redden on bruising
or become green in ammonia fumes. Mycol. Res. 94: 641-670.
Silva-Pinhati AC, Bacci Jr. M, Hinkle ML, Sogin M, Pagnocca FC. 2004. Low variation in ribosomal
DNA and internal transcribed spacers of the symbiotic fungi of leaf-cutting ants (Aftini:
Formicidae). Braz. J. Med. Biol. Res. 37: 1461-1472.
Singer R. 1986. The Agaricales in modern taxonomy. Ed. 4. Koeltz Scientic Books, Koenigstein. pp.
474-479,
Smith HV, Weber NS. 1987. Observations on Lepiota americana and some related species. Contr.
Univ. Mich. Herb. 16: 211-221.
Swofford DL. 2002. PAUP*: phylogenetic analysis using parsimony (*and other methods). Version
4.0b10. Sinauer Associates. Sunderland, USA.
Thompson JD, Gibson TJ, Plewniak F, Jeanmougin F, Higgins D. 1994. The CLUSTAL_X Windows
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Vellinga EC. 2000. Notes on Lepiota and Leucoagaricus. Type studies on Lepiota magnispora,
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ee
MYCOTAXON
Volume 106, pp. 379-383 October-December 2008
A new species of Dictyostelium
XTAO AN FIB SVU
'‘xidolanhe1121@yahoo.com.cn ? yuli966@126.com
Jilin Agricultural University
Changchun 130118, China
Abstract — A new species, Dictyostelium culliculosum, was isolated from forest soil
in China. Dictyostelium culliculosum is recognized by its special characteristics of
cell aggregations and sorophores and bases of sorophores. Detailed descriptions and
photographs of important life cycle stages are given based on the holotype.
Key words — dictyostelids, taxonomy
Introduction
The dictyostelids (cellular slime molds) have been known for more than a
century. They are often abundant in terrestrial ecosystems. The dictyostelids
have both animal-like (protozoan) and plant-like (fungal) characteristics. The
vegetative phase consist of small, independent amoeboid cells (myxamoebae)
that feed upon bacteria. They stream together in great numbers when the
available food becomes exhausted, and form pseudoplasmodia from which
subsequently develop sorocarps that consist of stalks and sori. The order
Dictyosteliales includes four genera, namely Dictyostelium, Polysphondylium,
Coenonia and Acytostelium (Kirk et al. 2001). The genus Dictyostelium was
described in 1869 (Raper 1984).
Materials and methods
Fifty samples for isolation of dictyostelids were collected during September 28
to September 29, 2007 in Jilin Province, and deposited under 4 °C condition.
The isolation method described by Cavender & Raper (1965) was adopted and
improved in this study. Five g of each sample was mixed with 50ml distilled
water, and the mixture was shaken for 1-2 h to suspend the cells of dictyostelids.
Then 0.5 ml of this suspension and approximately 0.4 ml of a heavy suspension
*Corresponding author
380 ... He & Li
of Escherichia coli were added to each of five 95 x 15 mm culture plates prepared
with hay infusion agar. Plates were incubated at 23 °C. Each plate was examined
once a day following 2 days incubation and then daily for up to 7 days. Isolates
were purified and subcultured for taxonomic examination on non-nutrient
agar using E. coli as the food source. Observations of early aggregations and
fruiting bodies were made after 24h of incubation at 23 °C. Sorocarps, stalks
and spores were measured, photographed using a Nikon dissecting microscope
(SMZ1500) and a Nikon light microscope (SMZ1000).
Taxonomy
Dictyostelium culliculosum Yu Li & Xiao-Lan He, sp. nov. Ficures1, 2
MycoBank MB 512298
Sorocarpia in agaro non nutricio cum E. coli ad 23 °C plerumque solitaria vel laxe
fasciculata, primo erecta vel semi-erecta aut decumbentia, interdum ramosa, rami longi
dispersi irregularesque, plerumque 1.2-5.0 mm longa. Sorophora plerumque sinuosa
inaequalia, typice e strato cellularum composita vel 2-3-cellulae, segmento terminali leviter
caesio vel diaphano, inconspicue angustata e basi ad apicem. Bases conspicuae, geniculatae,
multicellulares. Rami sorophorum mediorum superiorumque plerumque e_ strato
cellularum singulo compositi, basibus inconspicuis, plerumque ad angulum rectangulatum
ab axe principali abeuntes. Sori globosi vel citriformi, hyalino-albi, plerumque 50-100 um
in diametro. Sporae ellipticae vel oblongae, hyalini, sine polaribus granulis, plerumque
6.2-7.0 x 3.4-4.0 um. Cellularum aggregationes acervataeque sine rivulis, raro stellatae,
producentes unam vel aliquot sorogenam. Sorogena caesia vel schistacea, fortasse sorocarpia
formantia in situ vel post motum in distantia brevi. Myxamoebae parvae circa 6-14 x
8-16 um. Macrocystae globosae, numerus limitatus, plerumque 15-20 um in diametro;
microcystae globosae, numerosae, plerumque 5-9 um in diametro.
EryMo.oey: Latin, culliculosum, referring to the rounded mound-like aggregations.
Holotype: MRO008. Isolated from forest soil, Wangging, Jilin Province. Deposited at
the Herbarium of the Mycological Institute of Jilin Agricultural University (HMJAU),
Changchun.
When cultured at 23 °C on non-nutrient agar with E. coli, sorocarps solitary to
loosely clustered, erect or semi-erect at first, commonly 1.2-5.0 mm, usually
unbranched or with branches limited and spaced irregularly. Sorophores
inconspicuously tapering from bases to tips, some typically consist of one tier
of cells, and the others are 2-3-celled. Terminal segments of sorophores slightly
lavender or translucent. Bases conspicuous, knee-jointed. Branches of middle
to upper sorophores generally of one tier of cells, with inconspicuous bases and
mostly at a right angle with respect to the main axis. Sori white or colorless,
globose or citriform, commonly 50-100 um in diameter. Spores hyaline,
elliptical to oblong, commonly 6.2-7.0 x 3.4-4.0 um, without polar granules.
Cell aggregations typically developing as rounded mounds, usually without
stream formation, rarely form star-like shape and with inconspicuous streams
at later stage, producing one to several sorogens. Sorogens slightly lavender or
Dictyostelium culliculosum sp. nov. (China) ... 381
Figure 1. Dictyostelium culliculosum.
A, Sorocarps; B, Aggregation; C, Migrating pseudoplasmodium; D, E, Early (D) and late (E)
| sorogens; F, Sorophores and knee-jointed bases; G, Spores without granules; H, Microcyst;
| I, Macrocyst. Scale bars: A = 1 mm; B, C = 0.3 mm; D = 0.5 mm; E = 0.5 mm; F = 15 um; G =7
ens = 5 wm; 1 = 15 um.
blue grayish, may fruit in situ or migrate for some distance before sorocarp
_ formation is attempted. Myxamoebae small, about 6-14 x 8-16 um. Microcysts
| commonly formed by unaggregated myxamoebae, sometimes dominant in the
culture, globose, 5-9 um in diameter; macrocyst limited in number commonly
| 15-20 um in diameter.
| ComMENTs: This species can be distinguished from other species of
Dictyostelium by the combination of the following characteristics: (1) mound-
382 ... He & Li
FiGurRE 2. Dictyostelium culliculosum.
A, Sorocarps; B, Aggregation; C, Branch at a right angle with respect to the main axis;
D, Multicellular sorophore and knee-jointed base; E, 1-celled sorophore and knee-jointed base;
F, Elliptical spores.
like cell aggregations without streams; (2) terminal segments of sorophores
slightly lavender; (3) sorogens slightly lavender or blue grayish; (4) bases of
sorophores knee-jointed; (5) sorophores consist of 2-3 tiers of cells or typical
one tier of cells with knee-jointed bases; (6) sorocarps occasionally branched,
lateral branches consist of one tier of cells and mostly at a right angle in respect
to the main axis.
Sorocarps of Dictyostelium culliculosum closely resemble those of
D. brefeldianum H. Hagiwara (Hagiwara 1984) and D. citrinum Vadell, M.T.
Holmes & Cavender (Vadell et al. 1995). However, D. culliculosum sorophores
lack the capitate tips found in the latter two species. Furthermore, the new
species produces mound-like aggregations unlike the radiate aggregations
found in D. brefeldianum and D. citrinum. Lastly, almost all senescent mature
sorocarps of D. culliculosum lie prostrate on the substrate.
Acknowledgements
We express our deep appreciation to Profs. Jian-Yun Zhuang, AJS Whalley, and Tom
Hsiang for their valuable suggestions and kind help.
Dictyostelium culliculosum sp. nov. (China) ... 383
Literature cited
Cavender JC, Raper KB. 1965. The Acrasieae in nature. I. Isolation. American Journal of Botany
52: 294-296.
Hagiwara H. 1984. Review of Dictyostelium mucoroides Brefeld and D. sphaerocephalum (Oud.)
Sacc. et March. Bull. Natn. Sci. Mus., Tokyo, Ser. B. 10(1): 27-41.
Kirk PM, Cannon PE, David JC, Stalpers JA. (eds.) 2001. Ainsworth & Bisby’s Dictionary of Fungi
(ninth edition) prepared by CABI Bioscience. CABI International.
Raper KB. 1984. The Dictyostelids. Princeton University Press.
Vadell EM, Holmes MT, Cavender JC. 1995. Dictyostelium citrinum, D. medusoides and
D. granulophorum: three new members of the Dictyosteliaceae from forest soils of Tikal,
Guatemala. Mycologia 87(4): 551-559.
MYCOTAXON
Volume 106, pp. 385-398 October-December 2008
Records and two new species of Volvariella
(Pluteaceae, Agaricales) from Brazil
NELSON MENOLLI JR. & MARINA CAPELARI
menollijr@yahoo.com.br
Instituto de Botanica, Secao de Micologia e Liquenologia
Caixa Postal 3005, 01061-970 Sao Paulo, SP, Brazil
Abstract — Two new species of Volvariella, V. heterospora and V. nullicystidiata,
are described and illustrated from specimens collected at the Parque Estadual da
Cantareira, Sao Paulo, SP, Brazil. V. heterospora is characterized by its two types of
basidiospores, whereas V. nullicystidiata lacks cystidia. Volvariella bombycina and V.
perciliata, both collected in the same area, are also described, illustrated and recorded.
The latter represents the first record for Sao Paulo State and the second record for
Brazil. In addition, the specimens of Volvariella deposited at herbaria SP and collected
in remnants of the Atlantic forest of the metropolitan region of Sao Paulo, SP, Brazil,
were re-evaluated.
Key words — Basidiomycota, diversity, taxonomy
Introduction
_ Volvariella Speg. is a widely distributed genus whose species have been reported
from tropical, subtropical, and temperate regions of both eastern and western
hemispheres (Shaffer 1957). The genus is characterized by bulbangiocarpic
development with a membranous or fleshy volva, annulus absent, free and
pinkish lamellae, inamyloid basidiospores with a stramineous thickened wall,
| and bilateral convergent hymenophoral trama (Pegler 1983).
The genus comprises ca. 50 species (Kirk et al. 2001) with 17 mentioned for
Brazil. Montagne (1856) was the first author to record species of Volvariella
from Brazil, with V. cnemidophora (Mont.) Singer (as Agaricus cnemidophorus).
Rick (1930, 1961) recorded eight species of Volvariella for Rio Grande do Sul
State, viz. V. bombycina (as Volvaria bombycina), V. cnemidophora (as Volvaria
cnemidophora), V. fibrillosa (Bres. ex Rick) Singer (as Volvaria fibrillosa),
_V. pusilla (Pers.) Singer (as Volvaria parvula), V. surrecta (Knapp) Singer (as
| Volvaria loveana), V. taylorii (Berk. & Broome) Singer (as Volvaria taylorii),
|
386 ... Menolli & Capelari |
|
V. thwaitesii (Hook. f.) G.C. Rath (as Volvaria thwaitesii), and V. volvacea (Bull.)
Singer (as Volvaria volvacea). .
For Pernambuco State, Batista (1957) described V. oswaldoi (Bat.) Putzke |
(as Volvaria oswaldoi) and recorded V. gloiocephala (DC.) Boekhout & Enderle
(as Volvaria speciosa), and later Batista & Bezerra (1960) also mentioned |
V. cnemidophora. Singer (1989) described two new species, V. macrospora Singer !
and V. rondoniensis Singer, from Amazonas and Rondonia States respectively. |
For Sao Paulo State, V. bakeri (Murrill) Shaffer (Grandi et al. 1984, Pegler 1997),
V. bombycina (Bononi et al. 1981) and V. taylorii (Pegler 1997) were mentioned. |
For Parana State, ten species of Volvariella were mentioned by Meijer (2001, |
2006), including V. bombycina, V. earlei (Murrill) Shaffer, V. gloiocephala, |
V. hypopithys (Fr.) M.M. Moser, V. cf. murinella (Quél.) M.M. Moser,
V. perciliata, V. cf. pusilla var. taylorii (Berk. & Broome) Boekhout, |
V. rondoniensis, V. cf. striata N.C. Pathak and V. volvacea. |
Beside these species, there is the record of three species of Volvaria (Fr.) P. |
Kumm. for Rio Grande do Sul State that were not yet combined in Volvariella, |
viz. Volvaria pubipes (Peck) Sacc. (Rick 1938, 1961), Volvaria rhodomelas (Lasch) |
P. Kumm. (Rick 1930, 1938, 1961) and Volvaria umbonata var. brasiliensis Rick |
(Rick 1930). |
In this article, we report on four species of Volvariella recently collected |
at the Parque Estadual da Cantareira, Sao Paulo, SP, Brazil, two of which are
described as new. In addition, the specimens deposited as Volvariella at the |
Herbario do Estado Maria Eneyda P. Kauffmann Fidalgo (SP) and collected in |
remnants of Atlantic forest of metropolitan region of Sao Paulo City, SP, Brazil |
were also re-evaluated.
Material and methods
The specimens were collected at the Parque Estadual da Cantareira, a remnant |
of Atlantic forest in Sao Paulo State, Brazil and deposited at herbarium SP. The |
macroscopical descriptions and illustrations of basidiomata were based on |
fresh material. Color terms are from Kiippers (1979). The material deposited
as Volvariella at herbaria SP was revised from herbarium material originally |
collected at the Parque Estadual das Fontes do Ipiranga (PEFI), a remnant of |
Atlantic forest inside of urban area of Sao Paulo City, SP, Brazil. For microscopic |
analysis, the dried material was rehydrated in 70% ethanol followed by 5% ~
KOH or Melzer’s reagent. All microscopic illustrations were made with the aid
of a drawing tube. The notation “[a/b/c]” at the beginning of a set of spore |
data is to be read, “a spores were measured from ‘b’ basidiomata taken from
‘¢ collections’, and the Q represents the range of length/width quotients for all |
spores measured. The SEM images followed Baroni (1981).
|
Volvariella spp. nov. (Brazil) ... 387
DNA sequences of the large subunit (LSU) and internal transcribed space (ITS)
region of nuclear ribosomal DNA were obtained from three species, for future
phylogenetic molecular studies. GenBank accession numbers are cited below
the species name at the beginning of the formal description.
Results
Three collections deposited as Volvariella at SP and collected in Atlantic forest
remnants in the metropolitan area of Sao Paulo City were re-evaluated. Two
collections (SP47345 and SP47623) previously identified as V. bombycina were
destroyed by insects and moulds so that it was impossible to confirm their
identifications. Re-examination of the third collection labeled as V. bakeri
(SP128034) demonstrated that it represents a species of Pluteus Fr. section
Pluteus as indicated by the absence of a volva in the basidioma and presence
of horned metuloid cystidia, which is typical of section Pluteus. This result
invalidated the record of V. bakeri for Brazil as mentioned by Grandi et al.
(1984) and Pegler (1997).
_ The other specimens studied in this work collected at the Parque Estadual da
Cantareira are described below.
| Volvariella bombycina (Schaeff.) Singer FIGS. 1, 5A-B
GENBANK EU920669, EU920673
_MacrocHARACTERS — PILeus large, 140 mm diam., campanulate to convex,
_ slightly umbonate, whitish-cream, surface dry, smooth at center with yellowish
| fibrils toward the margin. LAMELLAE remote, pinkish, crowded, with lamellulae.
Stree 145 x 11(apex)-17(base) mm, yellowish-cream, tapering toward the
apex and slightly bulbous, flexuous, slightly fibrillose at the apex and slightly
| longitudinally striate toward the base. Votva 40 mm long, saccate, yellowish-
white, ample, well developed, often deliquescent, with scanty mycelium at the
base.
| MICROCHARACTERS — BASIDIOSPORES [20/1/1] 8.7(-10) x 6.2(-7.5) um
| (Q = 1.33-1.4), ellipsoid, smooth, slightly pinkish, inamyloid, thick-walled,
| guttulate, often 2-guttulate. Bastp1A (18.7—)20-23(-25) x (6.2-)7.5-8.7 um,
clavate, thin-walled, with small guttulae, 4-spored, with short sterigmata.
| PLEUROCYSTIDIA (36-)40-64(-70) x (7.5—)8.7-15(-17.5) um, appendiculate
or ventricose, sometimes strangulate at the apex or throughout extension,
fairly abundant and well distributed, thin-walled. CHEILocystip1A (50-)60-
94(-100) x (13.7-)16.2-24 um, clavate to slightly fusoid, very abundant,
thin-walled. LAMELLA TRAMA bilateral convergent, 50-89 um width, of thin-
| walled hyphae, 3.7-11.2 um diam., hyaline, sometimes pale yellowish, septate.
| SUBHYMENIUM cellular. PILEUS CONTEXT undifferentiated, composed by thin-
388 ... Menolli & Capelari
/
|
/
\
\
4
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Figure 1: Volvariella bombycina. a. Basidioma (Scale bar = 1 cm).
b. Basidiospores. c. Basidia. d. Pleurocystidia. e. Cheilocystidia. Scale bars = 10 um.
Volvariella spp. nov. (Brazil) ... 389
walled hyphae, 6.2-12.5 um diam., hyaline, septate. PILEIPELLIS composed by
thin-walled hyphae, 5-18 um diam., with yellowish content, arranged more or
less parallel to surface. VOLVAL ELEMENTS formed by thin-walled hyphae, 3.7-
13.7 um diam., hyaline or with light brown content, septate, well compacted
and parallel. CLAMP CONNECTIONS absent in all parts examined.
HABITAT AND SUBSTRATE — Solitary on decaying wood.
SPECIMENS EXAMINED — BRAZIL. SAO PAULO STATE: Sao Paulo, PARQUE ESTADUAL
DA CANTAREIRA, NUCLEO ENGORDADOR — 30.X.2007, Menolli Jr. et al. 151
(SP).
CoMMENTs — Volvariella bombycina is a widely distributed species and
recognized by its robust habit, pale silky-scaly pileus, well-developed saccate
volva, habitat on wood, and medium-sized basidiospores. Previous reports of
V. bombycina from Brazil include Rick (1961) for Rio Grande do Sul State, and
in the checklists of Bononi et al. (1981) and Meijer (2001, 2006) from Sao Paulo
and Parana States respectively.
Volvariella heterospora Menolli & Capelari, sp. nov. FIGs. 2, 5C-D
MycoBAnk MB 512149
Pileus 43 mm latus, conicus-convexus, umbonatus, atrocinereus, fibrillosus. Lamellae
liberae, roseae. Stipes 68 mm altus, apice usque ad 4 mm latus, basi 8 mm latis, albidus.
Volva 170 x 8 mm, adnata vel limbata, alba, externa coacta. Basidiosporae dimorphae:
typus I (6.4—)7.2(-8) x (4—)4.8(—5.6) um, late ellipsoideae vel ellipsoideae, raro elongatus;
typus II (8.8—)9.6-12(-13.6) x (4.8—)5.6-6.4 um, elongatus, raro ellipsoideae, plerumque
deformans. Basidia (16.2-)20-21 x 6.2-8.7 um, clavate, 4-sporigera. Pleurocystidia
(32-)37-49 x (6.2-)7.5-10(-11.2) um, versiformis, appendiculate, ventricosa vel
lageniformis, numerosa. Cheilocystidia (26—)34-41(-49) x (5-)7.5-10(-11.2) um,
clavata vel ventricosa. Trama himenophoralis inverse bilateralis, hyaline. Cuticula pilei
trichodermate, cellulis terminalis elongates, inflates, erectis vel suberectis. Hyphae omnes
defibulatae.
TYPE SPECIMENS — BRAZIL. SAO PAULO STATE: SAo Paulo, PARQUE ESTADUAL DA
CANTAREIRA, NUCLEO ENGORDADOR — 12.XII.2006, F. Karstedt & M. Capelari
832 (HOLOTYPE-SP).
EryMo_ocy: the name refers to the different types of basidiospores.
| MacrocHarRactTERs — PiLeus medium-sized, 43 mm diam., conico-convex,
| umbonate, dark gray with slightly darker center, surface dry, covered by
radially arranged gray fibrils, margin entire, sometimes exposing the lamellae.
| LAMELLAE free to remote, pinkish, crowded, with lamellulae. Stipe 68 x
| A(apex)-8(base) mm, white, tapering towards the apex, slightly subbulbose,
surface somewhat pruinose. VOLVA 170 x 8 mm, adnate to limbate, white, well
adhered to the stipe, enclosing it like a sock, with a cleavage line at one side of
stipe and at another side forming an adhered extension like a limb above the
base, outer surface slightly felted.
390 ... Menolli & Capelari
MICROCHARACTERS — BASIDIOSPORES [60/1/1] (6.4-)7.2(-8) x (4-)4.8(—5.6)
um (Q = 1.28-1.8), broadly ellipsoid to ellipsoid, rarely elongate, or bigger
(8.8—)9.6-12(-13.6) x (4.8-)5.6-6.4 (Q = 1.5-2.43), elongate, rarely ellipsoid,
frequently deformed and inequilateral or with the proximal end more tapering
than the distal end, smooth, hyaline to slightly pinkish, inamyloid, thick-
walled, guttulate, sometimes 2-guttulate, rarely 3-guttulate or nonguttulate.
BASIDIA (16.2—)20-21 x 6.2-8.7 um, clavate, thin-walled, with small scattered
guttulae, 4-spored. PLEUROCYSTIDIA (32-)37-49 x (6.2—)7.5-10(-11.2) um,
versiform, appendiculate or ventricose, sometimes lageniform 54-62 x
20-22 um, abundant and sparse, very hyaline and hard to see in KOH, thin-
walled. CHEILOCYSTIDIA (26—)34-41(-49) x (5-)7.5-10(-11.2) um, clavate to
ventricose, not abundant, hyaline and also difficult to see in KOH, thin-walled.
LAMELLA EDGE heteromorphous. LAMELLA TRAMA bilateral convergent, 44-50
um width, of thin or slightly thick-walled hyphae, 2.5-17.5 um diam., hyaline,
septate. SUBHYMENIUM cellular. PILEUS CONTEXT undifferentiated, 31-44 um
thick, composed by thin-walled hyphae, 3.7-18.7 um diam., hyaline, septate.
PILEIPELLIS trichodermic, thick, up to 640 um thick, composed by thin or
slightly thick-walled hyphae, 5-26 um diam., elongated, with terminal elements
slightly inflated, septate, with light brown content, arrangement erect or sub-
erect. VOLVAL ELEMENTS formed by thin-walled hyphae, 3.7-7.5 um diam.,
hyaline, frequently not septate, interwoven. CLAMP CONNECTIONS absent in all
parts examined.
HABITAT AND SUBSTRATE — Solitary on litter.
ComMMENTS — Volvariella heterospora is characterized by a small- to medium-
sized pileus that is conic-convex, umbonate, and gray with a slightly darker
center and by two types of basidiospores, one short, broadly ellipsoid to ellipsoid,
and the other bigger, elongate, rarely ellipsoid, and frequently deformed.
This species is similar to V. murinella, especially in macroscopic
characteristics (Boekhout 1986). But according to this author, V. murinella
has a low broad umbo, longer basidia [20-30(-35) x 7-10 um], much longer
cystidia [40-100 x 10-30(-50)], uniform basidiospores not exceeding 8.3 um
in length and 4.4 um in width [or 5 um according to Orton (1986)]. There
is no mention of the two basidiospore types or of the presence of bigger and
frequently deformed basidiospores, or of a volval extension along one side of
the stipe. Additionally, V. heterospora differs in pileus color and shape when
compared to the description of V. murinella by Lange (1935), who described
it as pale ashy gray and convex; Lange's illustration, which was also selected
by Boekhout (1986), represents a specimen with totally convex pileus, without
evident umbo or conic shape.
Volvariella spp. nov. (Brazil) ... 391
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Volvariella heterospora (holotype).
b. Small basidiospores. c. Large basidiospores. d. Basidia
FIGURE 2:
.
f. Cheilocystidia
e. Pleurocystidia.
Scale bars = 10 um.
392 ... Menolli & Capelari
Volvariella heterospora also resembles V. taylorii, but the latter is differentiated
by a gray, gray-olivaceous, hazel, brown, grayish avellaneous or grayish date-
brown volva (Lange 1935, Shaffer 1957, Pegler 1983, Orton 1986).
Volvariella nullicystidiata Menolli & Capelari, sp. nov. FIGS. 3, 5E-F
MycoBank MB 512150; GENBANK EU920670, EU920671
Pileus 66 mm latus, convexus, umbonatus, brunneous, pallidus cinereus ad marginem.
Lamellae liberae, roseae. Stipes 108 mm altus, albidus, apice usque ad 4.4 mm latus,
basi usque ad 8 mm latis. Volva 28 x 200 mm, saccata, trilobata, cupuliformis, cremeus,
cum mycelium basali. Basidiosporae (7.5-)8.7 x (5-)6.2(-7.5) um, ellipsoideae, raro late
ellipsoideae vel ovoideae. Basidia (22—)25-26(-28) x (7.5—)8.7-10(-11.2) um, clavata,
4-sporigera. Pleurocystidia et cheilocystidia absentibus. Trama hymenophoralis inverse
bilateralis, hyalina. Cuticula pilei hyphis repentibus. Hyphae omnes defibulatae.
TYPE SPECIMENS — BRAZIL. SAo PAULO STATE: Sao Paulo, PARQUE ESTADUAL
DA CANTAREIRA, NUCLEO ENGORDADOR — 31.1.2008, FE Karstedt et al. 1055
(HOLOTYPE-SP).
EryMo.oey: the name refers to the total absence of cystidia. -
MACROCHARACTERS — PILEUS medium-sized, 66 mm diam., convex
umbonate, dark brown at center, discoloring radially towards the margin,
margin light gray, surface dry, covered by radially arranged brown-grayish
fibrils, margin entire, slightly sulcate. LAMELLAE free, pinkish, with lamellulae.
STIPE 108 x 4.4(apex)—8(base) mm, white-cream, light sordid grayish on base,
tapering towards the apex, flexuous, slightly longitudinally striate. VoLvA 28 x
20 mm, saccate, beige, cupuliform, firm, well developed, 3-lobate, with cottony |
mycelium at the base.
MICROCHARACTERS — BASIDIOSPORES [20/1/1] (7.5—)8.7 x (5—)6.2(-7.5)
um (Q = 1.16-1.5), ellipsoid rarely broadly ellipsoid, with the distal end wider
and broader than the proximal end (egg-shaped), smooth, slightly pinkish,
inamyloid, thick-walled, guttulate, with a small apiculus hard to note and
seemingly truncate. Basrp1a (22-)25-26(-28) x (7.5-)8.7-10(-11.2) um, ;
clavate, thin-walled, sometimes with small guttulae, 4-spored. PLEUROCYSTIDIA
and CHEILOCYSTIDIA absent. LAMELLA EDGE fertile. LAMELLA TRAMA bilateral
convergent, up to 31 um width, of thin or slightly thick-walled hyphae, 3.7-15 |
jum diam., hyaline, septate. PILEUS CONTEXT undifferentiated, approximately
94 um thick, composed by slightly thick-walled hyphae, 3.7-12.5 um diam.,
hyaline, septate. PILEIPELLIS up to 225 um thick, composed by thin-walled
hyphae, 10-30 um diam., with light brown content, septate, arrangement more
or less parallel to the surface. VOLVAL ELEMENTS composed of thin or slightly
thick-walled hyphae, 2.5-15(-27) um diam., some inflated, hyaline, septate,
interwoven. CLAMP CONNECTIONS absent in all parts examined.
HABITAT AND SUBSTRATE — Solitary on soil.
Volvariella spp. nov. (Brazil) ... 393
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Ficure 3: Volvariella nullicystidiata (holotype). a. basidioma (scale bar = 1 cm).
b. basidiospores. c. basidia. scale bars = 10 um.
394 ... Menolli & Capelari
ComMEnts — Volvariella nullicystidiata is characterized by the complete absence
of cystidia. Few species of Volvariella lack cystidia, among them are V. acystidiata
N.C. Pathak and V. jamaicensis (Murrill) Shaffer. Volvariella acystidiata is quite
different from V. nullicystidiata because it has basidiospores that are much
longer [14.2-17.1(-19.9) x 8.5-10] as described in the protologue (Pathak
1975). In addition, V. acystidiata has a narrower (25 mm) cream-colored viscid
pileus, white and fragile volva, and basidia that are slightly longer (26.4-37.2 x
14.4-15.6 um) as described by Mora (1985) from material collected in Mexico.
However, V. nullicystidiata is closely related to V. jamaicensis. The latter differs
by the gregarious habit, slightly narrower pileus (50 mm) with slightly granular
surface, shorter stipe (50 x 3-5 mm), and a rather delicate, narrow, avellaneous
and slightly shorter (10-20 mm) volva as described in the protologue (Murrill
1911). Besides, V. jamaicensis has shorter and much narrower basidiospores,
which were cited in the protologue as 5 x 3 um and in the type study by Shaffer
(1957) as 6.2-8 x 3.3-4.3 um with a larger Q value of 1.54-2.08.
Volvariella perciliata Courtec. FIGS. 4, 5G-H.
GENBANK EU920668, EU920671
MACROCHARACTERS — PILEus small, 16-17 mm diam., convex with mammilate
umbo, grayish-brown (N_,A,,M,,) with a darker center, discoloring radially
towards the margin, surface dry, with short fibrils radially arranged, margin
ciliate throughout its extension. LAMELLAE free, pinkish, with lamellulae. STIPE
30-50 x 1.4-2 mm, white, cylindric, slightly pruinose, mostly on apical half,
covered by many humidity drops throughout its surface. Votva 5 x 8 mm,
saccate, white to slight brownish, firm, like an egg shell, 4-lobate, pubescent.
MICROCHARACTERS — BASIDIOSPORES [40/2/2] (6.2—)7.5-8.7(-10) x (3.7-)
5(-6.2) um (Q = 1.4-2), ellipsoid to elongate, smooth, inamyloid, thick-walled,
guttulate, often 2-guttulate, with a small apiculus hard to note. Basrp1a (21-)
23-29(-31) x (6.2-)7.5-8.7(-10) tm, clavate, thin-walled, with small guttulae,
4-spored, with short sterigmata. PLEUROCYSTIDIA 47-65 x 6.2-12.5 um,
fusiform occasionally slightly ventricose, sparse, thin-walled. CHEILOCYSTIDIA
abundant, like the pleurocystidia. LAMELLA TRAMA bilateral convergent,
37-44 um width, of thin-walled hyphae, 2.5-15 um diam., hyaline, septate.
SUBHYMENIUM cellular. PILEUS CONTEXT undifferentiated, approximately
125-187 um thick, constituted by thin-walled hyphae, 3.7-13.7 um diam.,
hyaline, septate. PILEIPELLIS trichodermic, up to 187 um thick, composed
by thin-walled hyphae, 6.2-15 um diam., elongated with the sub-terminal
elements inflated, septate, with light brown content, arrangement erect or sub-
erect. VOLVAL ELEMENTS composed of slightly thick-walled hyphae, 3.7-12.5
um diam., some slightly inflated, with light brown content, septate, interwoven.
CLAMP CONNECTIONS absent in all parts examined.
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396 ... Menolli & Capelari |
Volvariella spp. nov. (Brazil) ... 397
HABITAT AND SUBSTRATE — Solitary on wood or litter.
SPECIMENS EXAMINED — BRAZIL. SAO PAULO STATE: Sao Paulo, PARQUE ESTADUAL
DA CANTAREIRA, NUCLEO ENGORDADOR — 20.III.2007, Menolli Jr. et al. 115
(SP); 31.1.2008, E Karstedt et al. 1051 (SP).
CoMMENTs — Volvariella perciliata was originally described from French
Guyana (Courtecuisse 1991) and is characterized by a small basidioma, a
convex grayish-brown pileus with a mammillate umbo and a ciliate margin
throughout its extension. The Brazilian specimen has basidiospores that are
slightly longer than those described in the protologue (7-8 x 4-4.5 um) and
pleurocystidia slightly shorter than the type (45-75 x 7-12 um), complying
in all other characteristics. Since its original description, this species has been
recorded for Parana State (Meijer 2006). This collection represents the first
record for Sao Paulo State and the second record for Brazil.
Acknowledgments
The authors thank Dr. Clark L. Ovrebo, University of Central Oklahoma, and Dr. Régis
Courtecuisse, University of Lille, for critical review of the manuscript; Dr. Shaun R.
Pennycook, Landcare Research, for nomenclature review; Dr. Jefferson Prado, Instituto
de Botanica, who kindly revised the Latin diagnoses; Fernanda Karstedt, Instituto de
Botanica, for collecting some specimens and assistance with SEM; Anderson Luis dos
Santos, Instituto de Botanica, for assistance with formatting the plates; Klei R. Sousa for
assistance with ink illustrations; the Conselho Nacional de Desenvolvimento Cientifico
e Tecnolégico (CNPq) for the support and grant to the first author; and the Fundacao
de Amparo a Pesquisa do Estado de Sao Paulo (FAPESP grant 04/04319-2) for financial
support.
Literature cited
Baroni TJ. 1981. A revision of the genus Rhodocybe Maire (Agaricales). Beihefte zur Nova Hedwigia
6711-194,
Batista AC. 1957. Alguns Agaricaceae saprofiticos de Pernambuco. Mycopathologia et Mycologia
Applicata 8: 127-134.
Batista AC, Bezerra JL. 1960. Basidiomycetes vulgares no Nordeste brasileiro. Publicacdes do
Instituto de Micologia da Universidade do Recife 294: 1-30.
Boekhout T. 1986. Notulae ad Floram Agaricinam Neerlandicam - XII: small, saprophytic
Volvariella species in the Netherlands. Persoonia 13: 197-211.
FiGurE 5: SEM images. a-b. Basidiospores of Volvariella bombycina. c-d. Basidiospores of
V. heterospora (black arrow: large basidiospores; white arrow: small basidiospores).
e-f. V. nullicystidiata. e. Surface of lamella showing the absence of cystidia. f. Basidiospores.
g-h. V. perciliata. g. Surface of lamella with numerous pleurocystidia. h. Basidiospores and
pleurocystidia.
398 ... Menolli & Capelari
Bononi VLR, Trufem SFB, Grandi RAP. 1981. Fungos macroscépicos do Parque Estadual das
Fontes do Ipiranga, Sao Paulo, Brasil, depositados no Herbario do Instituto de Botanica. Rickia
oe Mook
Courtecuisse R. 1991. Eléments pour un inventaire mycologique des environs du saut Pararé
(Arataye) et de L’Inselberg des Nouragues, (Guyane francaise) V. Pluteaceae. (Pluteales,
Basidiomycota). Cryptogamic Botany 2(3): 136-152.
Grandi RAP, Guzman G, Bononi VL. 1984. Adicdes as Agaricales (Basidiomycetes) do Parque
Estadual das Fontes do Ipiranga, Sao Paulo, SP, Brasil. Rickia 11: 27-33.
Kirk PM, Cannon PF, David JC, Stalpers JA. 2001. Ainsworth and Bisby’s Dictionary of the fungi,
9th Edition Wallingford, CAB International.
Kiippers H. 1979. Atlas de los colores. Editorial Blume, Barcelona.
Lange JE. 1935. Flora Agaricina Danica I. Copenhagen, Recato.
Meijer AAR de. 2001. Mycological work in the Brazilian State of Paranda. Nova Hedwigia 72:
1052152)
Meijer AAR de. 2006. Preliminary list of the macromycetes from the Brazilian State of Parana.
Boletim do Museu Botanico Municipal 68: 1-55.
Montagne JPFC. 1856. Septieéme centurie de plantes cellulaires nouvelles, tant indigenes
quexotiques. Annales des Sciences Naturélles, Botanique série 4, 5: 333-374.
Mora VM. 1985. Nuevos registros del genero Volvariella en Morelos. Revista Mexicana de Micologia.
1291-100;
Murrill WA. 1911. The Agaricaceae of Tropical North America. Mycologia 3(6): 271-282.
Orton PD. 1986. Pluteaceae: Pluteus & Volvariella. In: Henderson DM, Orton PD, Watling R. (eds.).
British Fungus Flora: Agarics and Boleti, vol. 4. Edinburgh, Royal Botanic Garden.
Pathak NC. 1975. New species of Volvariella from Central Africa. Bulletin du Jardin Botanique
Naturelle de Belgique 45: 195-196.
Pegler DN. 1983. Agaric flora of the Lesser Antilles. Kew Bulletin Additional Series 9: 1-668.
Pegler DN. 1997. The agarics of Sao Paulo. Kew, Royal Botanic Gardens.
Rick J. 1930. Contributio IV ad monographiam Agaricacearum Brasiliensium. Broteria 24:
97-118)
Rick J. 1938. Agarici riograndenses. Lilloa 3: 399-455.
Rick J. 1961. Basidiomycetes eubasidii in Rio Grande do Sul - Brasilia. 5. Agaricaceae. Iheringia
Série Botanica 8: 296-450.
Shaffer RL. 1957. Volvariella in North America. Mycologia 49: 545-579.
Singer R. 1989. New taxa and new combinations of Agaricales (Diagnoses Fungorum Novorum
Agaricalium IV). Fieldiana Botany 21: 1-133.
MY COTA XON
Volume 106, pp. 399-401 October-December 2008
A new species of Ceraceomyces (Basidiomycetes)
from India
G. S. DHINGRA & AVNEET P. SINGH*
dhingragurpaul@gmail.com
Department of Botany, Punjabi University
Patiala 147 002 India
“Department of Biology, S D College
Barnala 148 101 India
Abstract - A new corticioid species, Ceraceomyces bizonatus, is described from Manali
hills in Himachal Pradesh.
Key words - Deodar, isodiametric hyphae, Kullu
While exploring the deodar forest in the Kothi area of Manali hills in district
Kullu of Himachal Pradesh, India, the authors made a collection on a decaying
log of Cedrus deodara with ceraceous hymenial surface. After detailed
macroscopic and microscopic comparisons with descriptions of known species
of genus Ceraceomyces (Julich 1972, Eriksson & Ryvarden 1973, Rattan 1977,
Thind & Dhingra 1985, Hjortstam 1998, Larsson & Larsson 1998) we found it
to belong to this genus but representing a species of its own. Typical characters
for the genus are the ceraceous basidiocarps, clavate, 4-spored basidia with a
basal clamp and ellipsoid to somewhat subglobose basidiospores. On the basis
of absence of cystidia, it comes closest to C. microsporus. However, a two-zoned
subhymenium, with a lower zone of vertical hyphae and an upper peculiar zone
of short-celled almost isodiametric encrusted hyphae, suggests that it represents
a new species. A sample of the basidiocarp was sent to Prof. Nils Hallenberg,
University of Géteborg, Sweden, who supported the concept of a new species.
Ceraceomyces bizonatus Dhingra & Avneet P. Singh sp. nov. FIGS. 1- 5
MycoBank MB507417
Basidiocarpus resupinatus, laxe adnatus, effusus, ad 225 um crassus in sectionis,
membranaceus-ceraceus ad ceraceus; superficies hymenialis laevis ad paulo tuberculatus,
fulvus; systema hyphale monomiticum; hyphae ad 4.7 um latum; basidii 16.5-24 x 3.1-5.5
um, clavatum, 4-sterigmatum, fibuligerum ad basis; basidiosporii 2.5-3.3 x 1.8-2.2 um,
ellipsoideus ad paulo subglobosum.
400 ... Dhingra & Singh
09000909
10 ym
i
=
3
Figs.1-4 Microscopic structures from basidiocarp of Ceraceomyces bizonatus
1. Basidiospores; 2. basidia; 3. generative hyphae; 4. vertical section of the basidiocarp.
Fig. 5. Ceraceomyces bizonatus basidiocarp showing hymenial surface.
————
Ceraceomyces bizonatus sp. nov. (India) ... 401
Holotypus: Himachal Pradesh: Kullu, Manali, Kothi,, super truncum Cedrus deodara,
Avneet 3612 (PUN), Septembre 12, 2004.
EryMo.ocy: The epithet refers to the two-zoned subhymenium.
Basidiocarps resupinate, loosely adnate, effused, up to 225 um thick in section;
cracks appear on drying; hymenial surface smooth to somewhat tuberculate,
yellowish brown; margins indeterminately thinning. Hyphal system monomitic;
generative hyphae up to 3 um wide, branched septate, clamped, thin-walled;
basal hyphae running somewhat parallel to the substratum; subhymenium
characteristic in having two zones - lower zone of vertical hyphae without
encrustation followed by a peculiar zone of short-celled almost isodiametric,
encrusted hyphae. Basidia 16.5-24 x 3.1-5.5 um, clavate, 4-sterigmate, with
a basal clamp; sterigmata upto 3.7 um long. Basidiospores 2.5-3.3 x 1.8-2.2
um, ellipsoid to somewhat subglobose, smooth, thin-walled, inamyloid,
acyanophilous.
Acknowledgements
Authors thank Prof. Nils Hallenberg (Goteborg, Sweden) for valuable suggestions and
peer review; Dr. B. M. Sharma (Department of Plant Pathology, CRKHPKV, Palampur,
India) for the peer review; Dr. Shaun Pennycook, Curator, ICMP, Auckland, New
Zealand for nomenclature review; Head of Department of Botany, Punjabi University
Patiala for providing infrastructure; and UGC DRS-SAP - II for financial assistance.
Literature cited
Eriksson J Ryvarden L. 1973. Corticiaceae of North Europe - I. pp. 59-286. Oslo.
Hjortstam K. 1998. A checklist to genera and species of corticioid fungi (Basidiomycotina,
Aphyllophorales). Windahlia 23: 1-54.
Julich W. 1972. Monographie der Athelieae (Corticiaceae: Basidiomycetes) Willdenowia 7: 1-283.
Larsson KH Larsson E. 1998. A molecular perspective on Ceraceomyces sublaevis. Folia Cryptog.
Estoniuca Fasc. 33: 71-78.
Rattan SS. 1977. The Resupinate Aphyllophorales of the North Western Himalayas. Bibliotheca
Mycologica 60: 1-427.
Thind KS Dhingra GS. 1985. Thelephoroid fungi of the Eastern Himalayas — I. Res. Bull. (Sci.) Pb.
Univ. Chd. 36(III-IV): 165-174.
Volume 106, pp. 403-407 October-December 2008
Annulatascus apiculatus sp. nov.,
a new freshwater ascomycete from the
semi-arid Caatinga biome of Brazil
FLAVIA RODRIGUES BARBOSA
Luis FERNANDO PASCHOLATI GUSMAO
faurb10@yahoo.com.br & lgusmao@uefs. br
Departamento de Ciéncias Biologicas,Laboratério de Micologia
Universidade Estadual de Feira de Santana
BR116 KM03, 44031-460, Feira de Santana, BA, Brazil.
&
HuZEPA A. RAJA
CAROL A. SHEARER
raja@illinois.edu & carolshe@illinois.edu
Department of Plant Biology, University of Illinois
Room 265 Morrill Hall 505 S. Goodwin Avenue, 61801 Urbana, IL, USA.
Abstract - During an inventory of lignicolous fungi in freshwater habitats in
northeastern Brazil, an interesting ascomycete belonging to the genus Annulatascus
was found. This fungus differs morphologically from other species of Annulatascus and
is herein described as a new species, A. apiculatus. The new species is characterized by
globose, black, semi-immersed perithecial ascomata with stout black necks; cylindrical,
unitunicate asci that have a relatively large bipartite, refractive apical apparatus;
hyaline, 0-3 septate ascospores with short, cellular, hyaline, tapering, bipolar apiculi,
and surrounded by a narrow mucilaginous sheath. The new species differs from other
described Annulatascus species in ascospore dimensions and the presence of bipolar
apiculi.
Key words — Annulatascaceae, diversity, systematic, submerged wood, taxonomy
Introduction
Aquatic ascomycetes are ecologically important, ubiquitous microbial saprobes
in freshwater environments (Wong et al. 1998, Tsui & Hyde 2004, Shearer
et al. 2007). About 557 species have been reported from freshwater habitats
worldwide (Shearer et al. 2008, http://www. fungi. life.uiuc.edu). It is surprising,
A404 ... Barbosa & al.
therefore, that no species have been reported from Brazil. Absence of such
reports most likely reflects lack of collecting efforts rather than absence of these
fungi from Brazilian freshwater habitats. To learn more about the freshwater
mitosporic and meiosporic ascomycetes in Brazil, we initiated a study of the
fungi colonizing dead plant substrates in freshwater habitats in the Caatinga
biome in northeastern Brazil.
During this study we collected an undescribed ascomycete from submerged
wood in a small stream. This ascomycete strongly resembled species in
the Annulatascaceae (Sordariomycetes), particularly those in the genus
Annulatascus. The most distinctive characteristic of this family and genus is the
presence of a very large ascus apical ring. Currently, Annulatascus includes 14
species (Tsui et al. 2002). Two of these species, A. citriosporus J. Frohl. & K.D.
Hyde and A. licualae J. Frohl. & K.D. Hyde, were described from terrestrial
habitats (Frohlich & Hyde 2000), while most other species have been reported
only from freshwater habitats in temperate and tropical latitudes (Hyde & Wong
2000, Cai et al. 2002, Tsui et al. 2002, http://www.fungi.life.uiuc.edu.).
The Brazilian specimen is described and illustrated herein as a new species of
Annulatascus and is compared to other species in the genus.
Materials and methods
STuDY SITE. Collecting trips were made to the Caatinga biome in the Serra da
Jibdia, one of nine hygrophilous forests that occur in the semi-arid region in
northeast Brazil (Velloso et al. 2002). The vegetation of this area is similar to
that of the Atlantic rain forest and has been described previously (Barbosa et al.
2007, Marques et al. 2007). Streams are bordered by bryophytes, pteridophytes
and several vascular plants.
COLLECTION TECHNIQUES. Submerged woody debris was collected from lentic
habitats and an unnamed stream in the Serra da Jibdia. Samples of submerged
dead plant material were placed in plastic bags and returned to the laboratory.
The plant material was then incubated at 25° C in Petri dish moist chambers
stored within 50 L plastic boxes with 200 ml sterile water plus 2 ml glycerol.
Samples were examined over four weeks for the presence of microfungal
fruiting bodies.
SPECIMEN EXAMINATION. Fruiting structures were located on the substrates
with a dissecting microscope and removed to a glass slide where they were
crushed and mounted in polyvinyl alcohol-glycerol (8.0 g in 100 ml of water,
plus 5 ml of glycerol). Measurements were made of fixed material. Dry material
and permanent slides were deposited in Herbarium HUEFS.
Annulatascus apiculatus sp. nov. (Brazil) ... 405
Taxonomy
Annulatascus apiculatus E.R. Barbosa & Gusmio, sp. nov. FIGs. 1-9
MycoBank MB512121
AscoMaTA 400-550 x 240-410 um, semi-immersa, globosa, nigra, coriacea, aggregata,
ostiolata. CoLLA 100-250 x 100-180 wm, cylindrica, atro-brunnea, periphysata.
PERIDIUM 15-60 um latis. PARAPHYSES 2.5-6 um crassae, hyalinae, filiformes, septatae,
glabro-tunicatae, simplices. Asct 175-250 x 10-13 um, 8-spori, cylindrici, unitunicati,
persistenti, pedicellati apparato apicale 6-7.2 x 1.8-2.4 um. ASCOSPORAE 23-36.5 x 8.8-10
um, uniseriatae, fusiformes, hyalinae, 0-3 septatae, laevae, cum spinulis in ambibus
extremitatibus 0.7-1.2 um et vagina mucilaginosa circundantes.
HoLotyPe: HUEFS 134723. BRAZIL. BAunrA: Santa Terezinha, Serra da Jibdia, on
submerged wood from a stream, 19.11.2008, coll. FR Barbosa.
EryMo_ocy: Latin, apiculatus referring to the apiculus present at both ends of the
ascospores.
ASCOMATA on wood, 400-550 x 240-410 um, clustered, semi-immersed,
globose, black, coriaceous, ostiolate. NECK 100-250 x 100-180 um, cylindrical,
dark brown, periphysate. PERIDIUM 15-60 um wide, dark brown. PARAPHYSES
2.5-6 x 75-100 um, broad at the base and tapering towards the apex, hyaline,
septate, smooth-walled, simple. Ascr 175-250 x 10-13 um, 8-spored, cylindrical,
unitunicate, persistent, pedicellate, with a large bipartite, refractive apical ring,
6-7.2 x 1.8-2.4 um. ASCOSPORES 23-36.5 x 8.8-10 um, uniseriate, fusiform,
straight, hyaline, 0-3 septate, not constricted at septa, with smooth, short,
cellular, hyaline tapering, bipolar apiculi; apiculi 0.7-1.2 um high; ascospore
surrounded by a narrow mucilaginous sheath.
CoMMENTs: The presence of an apiculus at both ends of the ascospores of
A. apiculatus differentiates this species from all other species of Annulatascus.
The bipolar apiculi on ascospores in the new species is quite different from
the bipolar pad-like appendages on ascospores of A. fusiformis K.D. Hyde
& S.W. Wong (Hyde & Wong 2000). Among the non-appendaged species of
Annulatascus, A. apiculatus is morphologically most similar to A. velatisporus
K.D. Hyde and A. aquaticus W.H. Ho et al. (Hyde 1992, Ho et al. 1999).
However, A. velatisporus has larger asci (220-290 x 12-18 ttm) and longer
non-septate ascospores surrounded by a gelatinous sheath (26-42 um), while
A. aquaticus has smaller asci (150-175 um) and non-septate ascospores with
smaller dimensions (19-24 x 6-7 um). Annulatascus apiculatus is also similar
to Annulusmagnus triseptatus (S.W. Wong et al.) J. Campb. & Shearer in having
3-septate ascospores (Campbell & Shearer 2004). The two species differ,
however, in that the ascospores of A. triseptatus are almost always 3-septate,
while those of A. apiculatus are non-septate when young and may become 3-
septate when older, and ascospores of A. triseptatus are concave or flattened on
406 ... Barbosa & al.
one side while those of A. apiculatus are not. To our knowledge, this represents
the first report of a freshwater ascomycete from Brazil.
Acknowledgements
We are deeply indebted to Dr. Jinx Campbell (University of Southern Mississippi) and Dr.
Amy Rossman (USDA) for kindly reviewing the manuscript and for many suggestions
that greatly improved it. The authors FR Barbosa and LFP Gusmao thank the CNPq for
financial support and the “Programa de Pés-graduacao em Botanica — PPGBot/UEFS”.
Literature cited
Barbosa FR, Gusmao LFP, Castafieda-Ruiz RF, Marques MFO, Maia LC. 2007. Conidial fungi from
the semi-arid Caatinga biome of Brazil. New species Deightoniella rugosa & Diplocladiella
cornitumida with new records for the neotropics. Mycotaxon 102: 39-49.
Cai L, Lumyong P, Zhang K, Hyde KD. 2002. New species of Annulatascus and Saccardoella from
the Philippines. Mycotaxon 84: 255-263.
Campbell J, Shearer CA. 2004. Annulusmagnus and Ascitendus, two new genera in the
Annulatascaceae. Mycologia 96: 822-833.
Frohlich J, Hyde KD. 2000. Palm microfungi. Fungal Diversity Research Series 3. Hong Kong.
Ho WWH, Hyde KD, Hodgkiss IJ. 1999. Ultrastructure of Annulatascus aquaticus sp. nov., a
freshwater Ascomycete on submerged wood from Hong Kong. Fungal Diversity 2: 119-128.
Hyde KD. 1992. Tropical Australian freshwater fungi II. Annulatascus velatispora gen. et sp. nov.,
Annulatascus bipolaris sp. nov. and Nais aquatica sp. nov. Aust. Syst. Bot. 5: 117-124.
Hyde KD, Wong SW. 2000. Annulatascus fusiformis sp. nov., a new freshwater ascomycete from the
Philippines. Mycologia 92: 553-557.
Marques MFO, Barbosa FR, Gusmao LFP, Castafieda-Ruiz RF, Maia LC. 2007. Conidial fungi from
the semi-arid Caatinga biome of Brazil. Cubasina microspora sp. nov., a note on C. albofusca,
and some new records for South America. Mycotaxon 102: 17-23.
Shearer CA, Descals E, Kohlmeyer B, Kohlmeyer J, Marvanova L, Padgett D, Porter D, Raja HA,
Schmit JP, Thorton HA, Voglmayr H. 2007. Fungal biodiversity in aquatic habitats. Biodiversity
and Conservation 16: 49-67.
Shearer CA, Raja HA, Schmit JP. 2008. Freshwater ascomycetes and their anamorphs. URL: http://
www.fungi.life.uiuc.edu (access 10/jun/2008).
Tsui CKM, Hyde KD. 2004. Biodiversity of fungi on submerged wood in a stream and its estuary in
the Tai Ho Bay, Hong Kong. Fungal Diversity 15: 205-220.
Tsui CKM, Ranghoo VM, Hodgkiss IJ, Hyde KD. 2002. Three new species of Annulatascus
(Ascomycetes) from Hong Kong freshwater habitats. Mycoscience 43: 383-389.
Velloso AL, Sampaio EVSB, Pareyn FCC (Eds). 2002. Ecorregides propostas para o Bioma Caatinga.
Associa¢ao Plantas do Nordeste, Instituto de Conservacao Ambiental, The Nature Conservancy
do Brasil, Recife.
Wong SW, Hyde KD, Jones EBG. 1998. Annulatascaceae, a new ascomycete family from the tropics.
Systema Ascomycetum 16: 17-25.
Fics. 1-9. Annulatascus apiculatus (from the holotype). 1. Immersed ascomata. 2. Longitudinal
section of the ascoma. 3. Ascus with eight ascospores. 4. Older, hyaline septate ascospore in ascus.
5. Ascus apical ring (arrowed). 6-9. Ascospores with bipolar apiculi (arrowed).
Scale bars: 1= 0.5 mm, 2 = 100 um, 3 = 25 um, 4 = 15 um, 5-9 = 10 um.
Annulatascus apiculatus sp. nov. (Brazil) ... 407
=
MYCOTAXON
Volume 106, pp. 409-412 October-December 2008
Re-identification of the anamorph of Leuconectria grandis
WEN- YING ZHUANG & JING LUO
zhuangwy@im.ac.cn & luojing999@hotmail.com
Key Laboratory of Systematic Mycology and Lichenology Laboratory
Institute of Microbiology, Chinese Academy of Sciences
Beijing 100101, China
Abstract — The anamorph of Leuconectria grandis was previously reported as
Gliocephalotrichum cylindrosporum based on morphological characteristics.
Re-examination of the anamorph and DNA sequence analysis of ITS and f-tubulin
indicate that the anamorph of L. grandis is distinct from G. cylindrosporum and
represents an unnamed species of Gliocephalotrichum. Following the recommendation
of the current International Code of Botanical Nomenclature, we avoid creating a name
for the anamorph of L. grandis.
Keywords — correction, morphology, taxonomy
Introduction
When the leaf-inhabiting Leuconectria grandis (Nectriaceae, Hypocreales)
was established, its anamorph was described at the same time and treated as
Gliocephalotrichum cylindrosporum based on morphological characteristics.
Distinctions between the Gliocephalotrichum material from China (ex type
culture of L. grandis) and the original description of G. cylindrosporum from
Thailand (Wiley & Simmons 1971) were discussed and treated as infraspecific
variations (Zhuang et al. 2007). Following the recent work on Gliocephalotrichum
by Decock et al. (2006), it becomes necessary to clarify the taxonomic placement
of the anamorph of L. grandis based on a combination of morphology and ITS
and f-tubulin sequence analysis.
Material and methods
Material used in this study is listed in TABLE 1. DNA was extracted from the
ex-type culture of Leuconectria grandis. DNA fragments and primers were
selected and DNA amplified in accordance with procedures outlined in Decock
etal. (2006). PCR products were purified using 3S Spin PCR Product Purification
* Author for correspondence.
410 ... Zhuang & Luo
Kit (Shenergy Biocolor for Life Science Co., Ltd.), then sequenced using the
same PCR primers with an ABI 3730 DNA analyzer by SinoGenoMax, Ltd.
Sequences were analyzed using the package MEGA4.10 (Tamura et al. 2007).
The resulting Neighbor-joining tree was derived from analyses of combined
sequences of ITS and f-tubulin genes with Glionectria tenuis as outgroup.
TABLE 1. Material studied
TAXA GENBANK NUMBER
ITS B-TUBULIN
Calonectria pauciramosa C.L. Schoch & Crous AF210886 AY078121
Calonectria pyrochroa (Desm.) Sacc. AY078115 AY078120
Gliocephalotrichum bacillisporum Decock & Huret 1 DQ374408 DQ374413
Gliocephalotrichum bacillisporum 2 DQ374409 DQ374414
Gliocephalotrichum bulbilium J.J. Ellis & Hesselt. 1 DQ374406 DQ377830
Gliocephalotrichum bulbilium 2 DQ374407 DQ377829
Gliocephalotrichum cylindrosporum B.J. Wiley & E.G. Simmons 1 DQ366705 DQ377841
Gliocephalotrichum cylindrosporum 2 DQ366706 DQ377842
Gliocephalotrichum longibrachium Decock & Charue 1 DQ278422 DQ377835
Gliocephalotrichum longibrachium 2 DQ278421 DQ377836
Gliocephalotrichum microchlamydosporum (J.A. Mey.) DQ366700 DQ374411
B.J.Wiley & E.G. Simmons 1
Gliocephalotrichum microchlamydosporum 2 DQ366701 DQ374412
Gliocephalotrichum ohiense L.H. Huang & J.A. Schmitt DQ366707 DQ374415
Gliocephalotrichum simplex (J.A. Mey.) B.J. Wiley & E.G.Simmons 1 DQ366703 DQ377839
Gliocephalotrichum simplex 2 DQ366704 DQ377840
Glionectria tenuis Crous & C.L. Schoch AF220980 AF320194
Leuconectria grandis Y. Nong & W.Y. Zhuang EF121859 U984072
Results and discussion
Re-examination of the Leuconectria grandis anamorph indicates that it differs
from any known species of Gliocephalotrichum by combining the following
features: sterile arms that more or less directly subtend a conidiogenous
penicillus in groups of 1-3 with a single arm sometimes growing distantly at the
lower portion of conidiophore or occasionally present on both the upper and
basal portions; the surface of the sterile arms often covered by an amorphous
substance; and narrow-cylindrical conidia that average 8.5-13 x 1.2-1.5 um.
Although the L. grandis anamorph more closely resembles G. cylindrosporum
than any other Gliocephalotrichum species with respect to conidial shape
and size, the position of the sterile arms is different. Compared to all species
producing similarly placed sterile arms, the L. grandis anamorph produces
narrower, longer conidia that distinguish it (Decock et al. 2006). The anamorph
appears to represent an unnamed species of Gliocephalotrichum.
Leuconectria grandis anamorph re-determined ... 411
To help determine the taxonomic position of the fungus, ITS and B-tubulin
gene sequences obtained from the known species of Gliocephalotrichum and
L. grandis were analyzed, and a Neighbor-joining tree was constructed (Fic. 1).
Our molecular data indicate that 1) Gliocephalotrichum is a monophyletic genus
with a relatively high bootstrap support (92%); 2) a sister-group relationship
can be recognized within the genus that correlates with the positioning of the
sterile arms; 3) G. cylindrosporum and G. simplex (both producing sterile arms
arising at a distance from the conidiogenous penicillus) form one highly (100%)
supported group; 4) the remaining species (all producing sterile arms that
directly subtend the conidiogenous penicillus) belong to a moderately (73%)
supported second group; and 5) there is a very low (42%) bootstrap support
for a relationship between L. grandis and G. longibrachium. Our data clearly
indicate that the L. grandis anamorph is not conspecific with G. cylindrosporum
but rather represents a distinct and unnamed species of Gliocephalotrichum.
In accordance with the current International Code of Botanical Nomenclature
(McNeill et al. 2006: Rec. 59A.3), we do not propose a separate name for the
new Gliocephalotrichum species. The holomorph name Leuconectria grandis
adequately designates both teleomorph and anamorph.
400 ; Gliocephalotrichum bulbilium 1
50 G. bulbilium 2
100 | G. bacillisporum 1
& G. bacillisporum 2
G. ohiense
05
73 400 | G. microchlamydosporum 1
G. microchlamydosporum 2
Leuconectria grandis
92 400 | G. longibrachium 1
G. longibrachiurn 2
400 - G. simplex 1
100 G. simplex 2
400 ; G.cylindrosporum 7
G.cylindrosporum 2
100 Calonectnia pauciramosa
Se Fe C. pyrochroa
Glionectria tenua
ey ae
0.01
Fie. 1 Neighbor-joining tree based on sequence analysis of ITS and f-tubulin genes, showing the
| relationships among species of the anamorphic genus Gliocephalotrichum.
412 ... Zhuang & Luo
Acknowledgments
The authors express their deep thanks to Dr. Shaun Pennycook for nomenclature
consultation, Dr. S. Pennycook and Dr. Yei-Zeng Wang for serving as the pre-submission
reviewers, and Dr. Lorelei Norvell for editorial revision and corrections of language.
This project was supported by the National Natural Science Foundation of China (nos.
30470009, 30670055), Ministry of Science and Technology of China (Special Project for
Fundamental Research no. 2006FY120100) and State 863 Project (no. 2008AA02Z312)
to WYZ.
Literature cited
Decock C, Huret S, Charue P. 2006. Anamorphic fungi from French Guyana: two undescribed
Gliocephalotrichum species (Nectriaceae, Hypocreales). Mycologia 98: 488-498.
McNeill J, Barrie FR, Burdet HM, Demoulin V, Hawksworth DL, Marhold K, Nicolson DH,
Prado J, Silva PC, Skog JE, Wiersma JH, Turland NJ. 2006. International Code of Botanical
Nomenclature (Vienna Code). Regnum Vegetabile No. 146.
Tamura K, Dudley J, Nei M, Kumar S. 2007. MEGA4: Molecular Evolutionary Genetics Analysis
(MEGA) software version 4.0. Molecular Biology and Evolution 24: 1596-1599.
Wiley BJ, Simmons EG. 1971. Gliocephalotrichum, new combinations and a new species. Mycologia —
63: 575-585.
Zhuang WY, Nong Y, Luo J. 2007. New species and new Chinese records of Bionectriaceae and
Nectriaceae (Hypocreales, Ascomycetes) from Hubei, China. Fungal Diversity 24: 347-357.
MYCOTAXON
Volume 106, pp. 413-418 October-December 2008
New species and new Chinese records of Dothideomycetes
WEN- YING LI*? & WEN- YING ZHUANG !
zhuangwy@im.ac.cn & ndwyli2002@163.com
'Key Laboratory of Systematic Mycology and Lichenology Laboratory
Institute of Microbiology, Chinese Academy of Sciences, Beijing 100101, China
°Graduate University of Chinese Academy of Sciences, Beijing 100049, China
Abstract — Some previous and recent collections of bitunicate ascomycetes from China
were examined. Saccardoella psidiicola is described as a new species. Dothidotthia
ramulicola, Macrovalsaria megalospora, Phaeosphaeria eustomoides and Plowrightia
periclymeni are recorded for the first time from China.
Key words — morphology, taxonomy
Introduction
Patouillard (1886) recorded the first Dothideomycetes from China when he
_ described Leptosphaeria delavayi Pat. together with six other new fungal species
collected by Delavay from Yunnan Province. Later research was attributed to
and summarized by Teng (1963), Tai (1979) and Eriksson & Yue (1988). More
| recently, Wang et al. (1999) and Lu et al. (2000), as well Yuan & Zhao (1994),
_ Hsieh & Chen (1994, 1996), Hyde (1995), Hsieh et al. (1997) and Li & Zhuang
| (2007, 2008) provided additional records to the Chinese Dothideomycetes
_ mycobiota. Some interesting fungi were encountered during our work on the
_ group from tropical and subtropical areas of China. Among them, one new
| species was discovered and four taxa are recorded for the first time from
| China.
Material and methods
| Herbarium material and recent collections from Hainan, Yunnan and Zhejiang
| provinces were studied. Ascomata from substrate were rehydrated and sectioned
| ata thickness of 10-20 um with a freezing microtome (YD-1508A, Yidi Medical
| Instrument Co.). Measurements were taken from the sections and from squash
| mounts in lactophenol cotton blue solution. Photographs were taken using a
* Author for correspondence.
414 ... Li & Zhuang
digital camera (Canon G5) connected to a Zeiss Axioskop 2 Plus microscope.
The collections studied are deposited in the Mycological Herbarium, Chinese
Academy of Sciences (HMAS).
Taxonomy
New species
Saccardoella psidiicola W.Y. Zhuang, W.Y. Li & K.D. Hyde, sp. nov. FIGS 1-5
MycoBank MB 512350
Pseudotheciis subglobosis vel piriformebus, 200-365 um diam., 360-510 ym alt.; ascis
clavatis, 8-sporis, 85-124 x 16.5-22 um; ascosporis fusiformibus, 7-10-septatis, hyalinis,
2/—-43.5 X 5.3-7.8 Um.
Ho.ortyPe: China. Paak Shan, on dead branches of Psidium guajava associated with at
least four other ascomycetes, V-1919, Otto A. Reinking, HMAS 10708. :
EtymMo_ocy: The specific epithet refers to the substrate of the fungus. |
Ascomata pseudothecial, subcortical, immersed in a pulvinate stroma,
solitary to gregarious, carbonaceous, subspherical to pyriform, individual —
locules 200-365 um diam. and 360-510 um high; hamathecia of trabeculate _
anastomosing pseudoparaphyses ca 0.8—1 um wide; asci bitunicate, outer wall |
very thin, with a thickened apex and a small ocular chamber, apical apparatus —
absent or unobvious, broadly clavate, 8-spored, 85-124 x 16.5-22 um; |
ascospores fusiform, hyaline, surrounded by a thick layer of hyaline gel sheath, |
mostly 7—10-septate, not constricted at septa, 27—43.5 x 5.3—7.8 um excluding ©
the sheath, irregularly biseriate to overlapping, spore sheath 1.2—-2 um thick. |
Anamorph unknown.
Notes: Among the known species of the genus Saccardoella (Spegazzini
1879, Saccardo 1895, 1899, Riedl 1968, Malhotra & Mukerji 1978, Hyde 1992,
Mathiassen 1993, Barr 1994, Hsieh et al. 1997, Barr et al. 1998, Tsui et al. 1998,
Hyde et al. 2000, Fallah & Shearer 2001, Cai et al. 2002), S. mangrovei K.D. |
Hyde from intertidal mangrove wood is the most similar to the Chinese fungus _
in ascospore shape and septation, but it differs significantly in fruitbody size
(390-845 um diam., 380-585 um high), longer and narrower asci (154-216 —
x 8.5-14 um) with an apical ring, and shorter spores (26-33 x 6-8 Um) —
diagonally uniseriate in ascus. S. graeweana (Blomb.) M.E. Barr et al. possesses
a somewhat similar spore shape, but has narrower asci (75-120 x 8-10 um) |
with an apical ring, and narrower spores (30-40 x 3-5.5 um) with 10-14 septa |
The fungus on Psidium guajava is new to the genus. i
The taxonomic position of Saccardoella remains unclear (Eriksson & |
Hawksworth 1991, Kirk et al. 2001). Because of its thickened ascus apex, |
small ocular chamber at apical portion of ascus, and trabeculate anastomosing ©
pseudoparaphyses among asci observed in the material, we treat the fungus as |
a bitunicate ascomycete.
ee
Saccardoella psidiicola sp. nov. & other Dothideomycetes (China) ... 415
prerrieninnnennnt
50 micron
‘5 40 micron
Fics 1-5. Saccardoella psidiicola (holotype): 1. Structure of ascocarp; 2. Hamathecium, showing
clavate asci associated with trabeculate pseudoparaphyses; 3, 4. Thickening at ascus apex;
5. Overlapping ascospores in ascus and an ascospore surrounded by a hyaline gel sheath.
New records for China
_ Dothidotthia ramulicola (Peck) M.E. Barr, Mycotaxon 29: 503, 1987.
= Sphaeria ramulicola Peck, Ann. Rep. New York State Mus. 25: 104, 1872.
SPECIMEN EXAMINED: China. Zhejiang, Tianmushan, alt. 500 m, on dead twigs, 6-XI-
2005, W.Y. Li & J. Luo 6785, HMAS 178097.
416 ... Li & Zhuang
Notes: Barr (1989) provided a detailed description of this species. The Zhejiang
collection is identical with the North American material, especially the red-
brown, straight and uniseptate ascospores measuring 19.5-25 x 9.5-14 um.
Macrovalsaria megalospora (Mont.) Sivan., Trans. Brit. Mycol. Soc. 65: 400, 1975.
= Sphaeria megalospora Mont., Ann. Sci. Nat., Bot., 2e sér., 14: 324, 1840.
= Macrovalsaria leonensis (Deighton) Petr., Sydowia 15: 300, 1962.
SPECIMENS EXAMINED: China. Hainan, Sanya, alt. 300 m, on dead twigs, 21-XI-2006,
W.Y. Li 7441, 7443, 7447, 7511, HMAS 178153, 178152, 178149, 178150; Hainan,
Ledong, alt. 1100 m, on dead twigs, 22-XI-2006, W.Y. Li 7475, HMAS 178151.
Notes: The brown, uniseptate ascospores that are constricted at the septum
and the skull cap-like germ apparatus at one end are diagnostic features for
the fungus. The genus is monotypic and Macrovalsaria leonensis is a synonym
(Sivanesan 1975, Hyde et al. 2000).
Phaeosphaeria eustomoides (Sacc.) Shoemaker & C.E. Babc.,
Can. J. Bot. 67: 1526; 1989:
= Leptosphaeria eustomoides Sacc., Fungi Venet. Nov. Vel. Crit. 2: 319, 1875.
SPECIMEN EXAMINED: China. Yunnan, Wenshan, on leaves of Oryza sativa, 11-VIII-
1938, Q.H. Wang & S.Z. Zhao 6310, HMAS 02310 (filed under Metasphaeria albescens
Thiim.).
NoTrEs: Our examination indicates that the Yunnan collection is identical with |
the morphology of Phaeosphaeria eustomoides described by Shoemaker Ss
Babcock (1989) based on a collection on Andropogon ischaemum.
Plowrightia periclymeni (Fuckel) Sacc., Syll. Fung. 2: 637, 1833.
= Dothidea periclymeni Fuckel, Jahrb. Nassauischen Vereins Naturk. 23-24: 223, 1870.
SPECIMEN EXAMINED: China. Zhejiang, Tianmushan, 1400 m, on small dead twigs of a
shrub, 2-XI-2005, W.Y. Li & J. Luo 6695, HMAS 178096.
DIAGNOSTIC FEATURES: Ascostromata initially immersed, becoming erumpent |
through the epidermis, pulvinate, with 2-3 locules, 0.5-0.65 x 0.16-017 mm;
pseudothecia subglobose, dark brown to nearly black, surface smooth or slightly
roughened, 96-226 um diam., 63-99 um high; peridium of textura angularis, |
composed of dark brown, thick-walled cells; asci bitunicate, clavate, 8-spored, |
66-96 x 11-14 um; ascospores fusiform, tapering towards both ends, 1-septate, —
slightly constricted at septum, hyaline, smooth-walled, irregularly biseriate, ©
13.5=16.5 %4.5~9.5 Um.
Notes: The pulvinate ascomata with small locules immersed, bitunicate asci, |
and hyaline ascospores with one septum indicate its position in the genus”
Plowrightia (Saccardo 1883, 1913, Arx & Miiller 1975, Barr 2001). The Chinese |
collection matches well the original description of Dothidea periclymeni on.
Lonicera periclymenum (Fuckel 1870, Saccardo 1833).
Saccardoella psidiicola sp. nov. & other Dothideomycetes (China) ... 417
Acknowledgments
The authors express their deep thanks to Prof. M. E. Barr, Prof. K. D. Hyde, Prof. D. H.
Pfister and Dr. G. Wade for taxonomic insights, providing useful references, and
consultation; Dr. L. Cai and Dr. E. H. C. McKenzie for serving as the pre-submission
reviewers and valuable suggestions; Dr. J. Luo for technical assistance; and Ms X.
Song for making sections. This project was supported by the National Natural Science
Foundation of China (nos. 30499340, 30670055) to WYZ.
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517-526.
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Barr ME. 2001. Revisionary studies on the Dothioraceae. Harvard Pap. Bot. 6: 25-34.
Barr ME, Holm L, Holm K. 1998. Reappraisal of Sphaeria ordinata sensu Broome. Mycotaxon 69:
31-34.
Cai L, Lumyong P, Zhang KQ, Hyde KD. 2002. New species of Annulatascus and Saccardoella from
the Philippines. Mycotaxon 84: 255-263.
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Ascomyc. 10: 27-67.
Eriksson OE, Yue J-Z. 1988. The Pyrenomycetes of China, an Annotated Checklist. University of
Umea, Sweden. 88 pp.
Fallah PM, Shearer CA. 2001. Freshwater ascomycetes: new or noteworthy species from north
temperate lakes in Wisconsin. Mycologia 93: 566-602.
Fuckel L. 1870 [1869-70]. Symbolae Mycologicae. Jahrb. Nassauischen Vereins Naturk. 23-24:
1-459.
Hsieh WH, Chen CY. 1994. Sivanesania, a new botryosphaeriaceous ascomycete genus on Rubus
from Taiwan. Mycol. Res. 98: 44-46.
Hsieh WH, Chen CY. 1996. Validation of Sivanesania. Mycol. Res. 100: 1106.
_ Hsieh WH, Chen CY, Sivanesan A. 1997. Some new ascomycetes from Taiwan. Mycol. Res. 101:
897-907.
| Hyde KD. 1992. The genus Saccardoella on intertidal mangrove wood. Mycologia 84: 803-810.
Hyde KD. 1995. Fungi from palms. XX. The genus Guignardia. Sydowia 47: 180-198.
Hyde KD, Taylor JE, Frohlich J. 2000. Genera of Ascomycetes from Palms. Fungal Diversity Press.
Hong Kong. 247 pp.
Kirk PM, Cannon PF, David JC, Stalpers JA. 2001. Ainsworth & Bisby’s Dictionary of the Fungi, 9th
Edition. CAB International, Wallingford. 655 pp.
| Li W-Y, Zhuang W-Y. 2007. Re-examinations of Botryosphaeriaceae (Dothideomycetes) on deposit
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Lu BS, Hyde KD, Ho WH, Tsui KM, Taylor JE, Wong KM, Yanna, Zhou DQ. 2000. Checklist of
Hong Kong Fungi. Fungal Diversity Press, Hong Kong. 207 pp.
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Mathiassen G. 1993. Corticolous and lignicolous Pyrenomycetes s. lat. (Ascomycetes) on Salix along
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272-276.
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Sivanesan A. 1975. Redisposition and descriptions of some Amphisphaeria species and a note on
Macrovalsaria. Trans. Brit. Mycol. Soc. 65: 395-402.
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MYCOTAXON
Volume 106, pp. 419-422 October-December 2008
Corticioid fungi (Basidiomycota) from Madeira Island
M.T. TELLERIA’, I. MELO?, M. DUENAs’, I. SALCEDO},
J. CARDOSO’, J.L. RODRIGUEZ-ARMAS* & E. BELTRAN-TEJERA®
‘telleria@rjb.csic.es
‘Real Jardin Botanico, CSIC
Plaza de Murillo 2, 28014 Madrid, Spain
?Jardim Botanico (MNHN), Universidade de Lisboa / CBA-FCUL
Rua da Escola Politécnica 58, 1250-102 Lisboa, Portugal
*Dpto. de Biologia Vegetal y Ecologia (Botanica), Universidad del Pais Vasco
Aptdo. 644, 48080 Bilbao, Spain
“Dpto. de Biologia Vegetal (Botanica), Universidad de La Laguna
38071 La Laguna Tenerife, Islas Canarias, Spain
Abstract — The first catalogue of corticioid fungi from Madeira Island is presented. The
study covered 18 localities and 561 samples were analyzed. This catalogue includes 110
species, 89 of them new to the island. They belong to 50 genera, of which Hyphodontia,
Hyphoderma, Trechispora, Stereum, Botryobasidium and Sistotrema are the more
significant. The presence of Stereum pseudorimosum, Subulicystidium perlongisporum
and Tubulicium dussii on the island is also remarkable. The complete catalogue is
available at http://www.mycotaxon.com/resources/weblist.html.
Key words — Aphyllophorales, fungal diversity, lignicolous fungi, Macaronesia,
Portugal, species inventory
Introduction
Madeira Island, situated between 32°38’ and 32°52’ N latitude and 16°39’ and
17°16’ W longitude, is the largest and highest island of the Madeira archipelago
and has a surface of 742 km®. It is truly volcanic, of Tertiary origin, without
connection to any major land masses. Its relief is dominated by Pico Ruivo
(1861 m), which divides the island into a North and South part, and Pico do
Arieiro (1810 m); the landform is quite complex, characterized by deep ravines
and gorges. The coastline is defined by inaccessible high cliffs and abundant
caves. The climate includes two main types, mediterranean and temperate with
a sharp altitudinal and north-south climatic gradient, with an average annual
420 ... Telleria & al.
temperature ranges between 15 to 20 °C and the rainfall ranges from 250 to
750 mm.
The laurisilva is the Macaronesian subtropical mountain moist forest that
characterizes Madeira and Canary archipelagos (Jardim & Sequeira 2008).
The wildlife of Madeira has been severely affected by human activity. Native
timber was heavily exploited over a long period of time, beginning shortly after
colonization in the 15" century. The laurisilva, which used to cover most of the
Madeira Island, persists now as a relict forest covering 16% of the Island.
Only a few corticioid species have been reported from Madeira (Holl 1830,
Torrend 1909, 1912, 1913; Dennis et al. 1977, Dias & Lucas 1980, Calonge
& Sequeira 2003, Capelo et al. 2007, Duefias et al. 2008). These species are
specifically mentioned in the catalogue.
Material and methods
18 localities were surveyed over three periods (spring 1998, summer 2000 and
autumn 2006). All potential substrates, indigenous and introduced taxa were
examined. 561 samples were studied following classical methods: thin, freehand |
sections were mounted in KOH (5%) and/or Melzer reagent and examined |
under Olympus BH 50 and Olympus BX 50 microscopes. The specimens have |
been deposited in BIO, LISU, MA-Fungi, and TFCMic herbaria.
Results
This first catalogue for the corticioid fungi from Madeira Island (http://www.
mycotaxon.com/resources/weblist.html.), so far, includes 110 species, of which |
89 are new to this island. They represent 50 genera with Hyphodontia (10
species), Hyphoderma (7 species), Stereum (7 species), Trechispora (7 species), _
Botryobasidium (6 species), and Sistotrema (5 species) being the most significant. |
The following 89 species are new to Madeira: Aleurodiscus aurantius (Pers.)
J.Schrot., Amylostereum laevigatum (Fr.) Boidin, Amyloxenasma allantosporum
(Oberw.) Hjortstam & Ryvarden, Aphanobasidium filicinum (Bourdot) Jiilich,
A. sphaerosporum Boidin & Gilles, Athelia decipiens (Hohn. & Litsch.) J.Erikss., |
Athelopsis glaucina (Bourdot & Galzin) Oberw. ex Parmasto, Botryobasidium _
aureum Parmasto, B. candicans J.Erikss., B. conspersum J.Erikss., B. danicum
J.Erikss. & Hjortstam, B. subcoronatum (Hohn. & Litsch.) Donk, B. vagum
(Berk. & M.A.Curtis) D.P.Rogers, Brevicellicium olivascens (Bres.) K.H.Larss.
& Hjortstam, Byssomerulius corium (Pers.) Parmasto, Cabalodontia queletii
(Bourdot & Galzin) Piatek, C. subcretacea (Litsch.) Piatek, Ceraceomyces eludens
K.H.Larss., Ceratobasidium cornigerum (Bourdot) D.P.Rogers, Coniophora \
puteana(Schumach.) P.Karst., Cylindrobasidium evolvens (Fr.) Jiilich, Fibrodontia |
Corticioids from Madeira ... 421
gossypina Parmasto, Hymenochaete cinnamomea (Pers.) Bres., H. corrugata
(Fr.) Lév., Hyphoderma argillaceum (Bres.) Donk, H. definitum (H.S.Jacks.)
Donk, H. litschaueri (Burt) J.Erikss. & A.Strid, H. medioburiense (Burt) Donk,
H. occidentale (D.P.Rogers) Boidin & Gilles, H. roseocremeum (Bres.) Donk,
H. transiens (Bres.) Parmasto, Hyphodontia alutaria (Burt) J.Erikss., H. arguta
(Fr.) J.Erikss., H. aspera (Fr.) J.Erikss., H. bugellensis (Ces.) J.Erikss., H. crustosa
(Pers.) J.Erikss., H. erastiiSaaren. & Kotir., H. nespori(Bres.) J.Erikss. & Hjortstam,
H. rimosissima (Peck) Gilb., H. spathulata (Schrad.) Parmasto, Metulodontia
nivea (P.Karst.) Parmasto, Mycoacia nothofagi (G.Cunn.) Ryvarden, M. uda
(Fr.) Donk, Peniophora limitata (Chaillet) Cooke, P meridionalis Boidin,
P. pilatiana Pouzar & Svréek, Peniophorella praetermissa (P.Karst.) K.H.Larss.,
Phanerochaete martelliana (Bres.) J.Erikss. & Ryvarden, Ph. sordida (P.Karst.)
J.Erikss. & Ryvarden, Phlebia lilascens (Bourdot) J.Erikss. & Hjortstam,
Ph. livida (Pers.) Bres., Ph. rufa (Pers.) M.P.Christ., Ph. subserialis (Bourdot &
Galzin) Donk, Phlebiella ardosiaca (Bourdot & Galzin) K.H.Larss. & Hjortstam,
Ph. tulasnelloidea (Hohn. & Litsch.) Ginns &M.N.L.Lefebvre, Phlebiopsisravenelii
(Cooke) Hjortstam, Pteridomyces bananisporus Boidin & Gilles, Radulomyces
confluens (Fr.) M.P.Christ., Scopuloides hydnoides (Cooke & Massee) Hjortstam
& Ryvarden, Scytinostroma hemidichophyticum Pouzar, Sc. ochroleucum (Bres.
& Torrend) Donk, Sistotrema brinkmannii (Bres.) J.Erikss., S. diademiferum
(Bourdot & Galzin) Donk, S. hispanicum M.Duefas et al., S. octosporum
(J.Schrot. ex Hohn. & Litsch.) Hallenb., S. subtrigonospermum D.P.Rogers,
Sistotremastrum suecicum Litsch. ex J.Erikss., Sistotremella perpusilla Hjortstam,
Steccherinum fimbriatum (Pers.) J.Erikss., Stereum ochraceoflavum (Schwein.)
| Sacc., St. pseudorimosum Boidin & Gilles, St. rugosum Pers., Subulicystidium
_ longisporum (Pat.) Parmasto, S. perlongisporum Boidin & Gilles, Tomentellopsis
| echinospora (Ellis) Hjortstam, Trechispora alnicola (Bourdot & Galzin) Liberta,
_ Tr. cohaerens (Schwein.) Jiilich & Stalpers, Tr farinacea (Pers.) Liberta,
| Tr. microspora (P.Karst.) Liberta, Tr. minima K.H.Larss., Tr. praefocata (Bourdot
| & Galzin) Liberta, Tubulicium dussii (Pat.) Oberw. ex Julich, T. vermiferum
| (Bourdot) Oberw. ex Jiilich, Tubulicrinis accedens (Bourdot & Galzin) Donk,
__ T. calothrix (Pat.) Donk, T. chaetophorus (Héhn.) Donk, T. subulatus (Bourdot
| & Galzin) Donk, Xenasma pruinosum (Pat.) Donk, X. pulverulentum (Litsch.)
| Donk.
| Aremarkable feature is the presence in the island of Stereum pseudorimosum,
| Tubulicium dussii and Subulicystidium perlongisporum, species with a similar
_ disjunct geographical pattern.
Finally a Nomina incertae of four names recorded by Torrend (1909, 1912,
_ 1913) that require reevaluation was established.
422 ... Telleria & al.
Acknowledgments
Financial support was given by DGI project n° CGL-2005-01192/BOS. Our thanks are
due to Dr. Francisco D. Calonge and Dr. Kurt Hjortstam for pre-submission review of
the manuscript, as well as to Dr. Susana Fontinha, director of Parque Natural da Madeira,
for authorizing us to collect fungi in the archipelago and to Carlos Santos, Manuel José
and Paulo Moniz for their assistance in local travel.
Literature cited
Calonge FD, Sequeira MM. 2003. Contribucidén al catalogo de los hongos de Madeira (Portugal).
Boletin de la Sociedad Micolégica de Madrid 27: 277-308.
Capelo J, Sequeira MM, Jardim R, Mesquita S. 2007. Biologia e ecologia das florestas das ilhas.
2 Madeira. 81-134, in Silva JS (ed.), Acores e Madeira. A floresta das ilhas. Lisboa, Fundacao
Luso-Americana para 0 Desenvolvimento.
Dennis RGW, Reid DA, Spooner BM. 1977. The fungi of the Azores. Kew Bulletin 32(1): 85-136.
Dias MRS, Lucas MT 1980. Fungi of Madeira and Selvagem Islands. Boletim da Sociedade
Broteriana, 2.? Série 53: 469-476.
Duenas M, Telleria MT, Melo I, Rodriguez-Armas JL, Beltran E. 2008. A new species of
Candelabrochaete (Polyporales, Basidiomycota). Mycotaxon 103: 299-305.
Holl F. 1830. Verzeichniss der auf der Insel Madeira beobachteten Pflanzen, nebst Roce
einiger neuen Arten. Flora oder Botanische Zeitung (Regensburg). 13(24): 369-392.
Jardim R, Sequeira MM. 2008. The vascular plants (Pteridophyta and Spermatophyta) of the Madeira
and Selvagens archipelagos. 157-178, in PAV Borges et al. (eds.), A list of the terrestrial fungi,
flora and fauna of Madeira and Selvagens archipelagos. Funchal and Angra do Heroismo.
Torrend C. 1909. Premiére contribution pour étude des champignons de I’Ile de Madere. Brotéria,
Série Botanica 8: 126-144.
Torrend C. 1912. Deusiéme contribution pour létude des champignons de I’Ile de Madere. Brotéria,
Série Botanica 10: 29-49.
Torrend C. 1913. Troisieme contribution pour létude des champignons de I'lle de Madere. Brotéria,
Série Botanica 11: 165-181.
Volume 106, pp. 423-429 October-December 2008
Irpex hacksungjii sp. nov. (Polyporaceae)
from Korea
JIN SUNG LEE’, CHANGMU KIM? & YOUNG Woon Lim?
‘jinsung@me.go.kr
*snubull@me.go.kr
>youngwlim@me.go.kr
National Institute of Biological Resources
Incheon 404-708, Korea
Abstract — Irpex is one of the most common polypore genera and characterized by its
poroid to hydnaceous hymenophores, encrusted cystidia, and simple-septate hyphae.
An Irpex specimen with conspicuous morphological differences from I. lacteus and I.
hydnoides was collected in Korea and this taxon is proposed as a new species, Irpex
hacksungii.
Key words — Basidiomycota, taxonomy, phylogeny, ITS
Introduction
|. The genus Irpex Fr. (Polyporaceae, Basidiomycetes) is characterized by its poroid,
irpicoid, hydnaceous hymenophores, encrusted cystidia, and simple-septate
hyphae (Cejp 1931, Gilbertson & Ryvarden 1986, Ryvarden & Gilbertson
| 1993). Because of a broad generic concept, the genus Irpex included numerous
| species. However, Maas Geesteranus (1974) treated many Irpex species as
synonyms of I. /acteus (Fr.) Fr. and moved all other species from the genus Irpex
to other genera. The monotytic genus Irpex remained stable then for thirty
years (Ryvarden & Gilbertson 1993). Recently, Lim & Jung (2003) included
a second species, Irpex hydnoides Y.W. Lim & HS. Jung, in this monotypic
genus. I. hydnoides is distinguished from I. lacteus based on hymenophoral
configuration and cultural and molecular characters.
_ The national biological inventory organized by the National Institute of
Biological Resources (NIBR, www.nibr.go.kr) is currently underway and has
yielded numerous collections of fungi from Korea. During observations of dry
Specimens at NIBR, a previously unknown species of Irpex was encountered.
This fungus is distinct from other Irpex species based on morphological and
molecular characters. Therefore, it is described in this paper as a new species.
424 ... Lee, Kim & Lim
Materials and methods
Specimens and observations
Macroscopic and microscopic characters were based on voucher specimens
deposited at Seoul National University Fungi Herbarium (SFC) with duplicates
at NIBR. The measurements and drawings were made on slide preparations
mounted with 3% (w/v) potassium hydroxide and stained with 1% (w/v)
phloxine in water (Largent et al. 1977) using a Nikon microscope and drawing
tube. |
Molecular analyses
Specimens that used on molecular approaches are listed in TABLE 1. Total DNA
was extracted from dried specimens using a Genomic DNA Extraction Kit
(Bioneer, Korea). Extracted DNA was stored at —20°C. The internal transcribed
spacer (ITS) region was amplified using primers ITS5 and ITS4 (White et al.
1990). PCR amplification was performed as described by Lee et al. (2000).
Amplified products were visualized by electrophoresis on a 1% agarose gel,
and purified with the PCR purification kit (Bioneer, Korea). Sequencing was
performed using the primers described above on an ABI 3730XL automated
sequencer (Applied Biosystems, USA). The sequence was proofread, edited,
and aligned with other Irpex species ITS sequences retrieved from the GenBank
using the jPHYDIT program (Jeon et al. 2005). Phylogenetic trees were inferred
from sequence alignment using neighbor-joining (NJ), maximum parsimony
(MP) and maximum likelihood (ML) methods implemented in PAUP 4.0b10
(Swofford 2002). Likelihood settings were parameterized using the best-fit
model (TrN+I) selected by AIC in Modeltest 3.7 (Posada & Crandall 1998).
TABLE 1. Irpex specimens used in this study.
SPECIES VoucHER No. LOCALITY Aaya
I. hacksungii SNU m-990326-16* Korea EU301650
I. hydnoides SNU m-971215-19 Korea AF479667
I. hydnoides SNU m-971011-12 Korea AF479668
I. lacteus SNU m-951007-39 Korea AF479666
I. lacteus SNU m-971006-12 Korea AF163046
* SNU, Seoul National University Herbarium, Seoul, Korea, Holotype.
Irpex hacksungii sp. nov. (Korea) ... 425
FicurE 1. Basidiocarps (A) and microscopic features (B) of Irpex hacksungii (holotype).
a, basidiospores. b, basidia. c, lamprocystidia. d, generative hyphae. e, skeletal hyphae.
Scale bars: A = 5 mm, B = 10 um.
Results
Taxonomy
Irpex hacksungii J.S. Lee & Y.W. Lim, sp. nov. FIGURE 1
MycoBAnk MB 512279
Basidiocarpus annuus, resupinatus vel effuso-reflexus; superficies canus, tomentoso-
hirsuto; hymenium cremeo-lutea vel ochraceo, hydnoidea, 2 mm longis; systema hypharum
dimiticum; hyphae generatoriae tenuitunicatae, 2-3.5 um; lamprocystidia incrustata,
24-33 x 5-6.5 um; basidia clavata, 4-sterigmatibus, 26-32 x 4.5-5.7 um; basidiosporae
: ellipsoideae, hyalinae, laeves, non-amyloideae, 5.0-6.2 x 3.0-3.5 um.
Holotypus in herbarium SNU; numero accessiono m-990326-16.
: Erymo oey: “hacksungii’, named in honour of Dr. Hack Sung Jung, a pioneering Korean
: mycologist. Among his many notable contributions to mycological research in Korea
: were excellent descriptions and keys for identifying polyporoid genera and species.
Basidiocarps annual, usually resupinate or effused-reflexed; pilei protrude up
to 3 mm, upper surface light gray; hymenophore cream, ocher to gray, tooth
up to 2 mm long; margin loose and often somewhat upcurved. Hyphal system
dimitic; generative hyphae thin-walled with frequent branching, frequently
426 ... Lee, Kim & Lim
septate without clamp connection, 2-3.5 um wide; skeletal hyphae hyaline,
thick-walled, rarely simple septate, 5-6 um wide. Lamprocystidia abundant,
thick-walled, heavily incrusted at apex, 24-33 x 5-6.5 um, incrusted part 10-14
x 5-6.5 um. Basidia clavate, 4-sterigmata, 26-32 x 4.5-5.7 tm, simple-septate at
the base. Basidiospores elliptical, hyaline, smooth, negative in Melzer’s reagent,
5.0-6.2 x 3.0-3.5 um.
COLLECTIONS EXAMINED: KOREA: Mungyeongsaejae, Mungyeong-si, Gyeongsangbuk-
do, ona fallen branch of an unknown deciduous tree, 26 March 1999, Y. W. Lim, deposited
in Seoul National University Herbarium (SNU), Specimen No. SNU m-990326-16
(HOLOTYPE), and National Institute of Biologial Resources (NIBR), Specimen No.
NIBRFG0000001610 (ISOTYPE).
Notes — Although this fungus is macroscopically similar to Irpex hydnoides,
the teeth are shorter and similar to those of Steccherinum Gray species.
Microscopic differences of I. hacksungii include: smaller lamprocystidia than
those in I. hydnoides and I. lacteus (I. hacksungii: 24-33 x 5-6.5 um, I. hydnoides:
40-57 x 8-10 um, I. lacteus: 65-80 x 6.8-11.5 um); smaller basidiospores than
I. hydnoides (I. hacksungii: 5-—6.2 x 3-3.5 um, I. hydnoides: 5.9-6.7 x 3.5-4 um);
and smaller basidia than I. hydnoides (I. hacksungii: 26-32 x 4.5-5.7 um, I.
hydnoides: 30-32 x 5.5-6.5 um) (TABLE 2).
TABLE 2. Morphological characters of Irpex species.
SPECIES Irpex hacksungii Irpex hydnoides Irpex lacteus
Resupinate to effuse- Resupinate to effuse- Resupinate to effuse-
BASIDIOCARP
reflexed reflexed reflexed
ens hydnoid, hydnoid, poroid, irpicoid, hydnoid,
yellow to gray deep yellow milky white to cream
HyYPHAL SYSTEM dimitic dimitic dimitic
LAMPROCYSTIDIA (tm) 24-33 x 5-6.52 40-57 x 8-10 65-80 x 6.8-11.5
(encrusted part) (10-14 x 5-6.5) (23-25 x 8-10) (25=36-8= =>)
BASIDIA (um) 26-32 x 4.5-5.7 30-32 x 5.5-6.5 24-25 x 4-5
BASIDIOSPORES (tm) 5-6.2 x 3-3.5 5.9-6.7 X 3.5-4 5.5-6 xX 2.2-2.8
* Twenty (20) cystidia, basidia and spores of each specimen were measured.
Sequence analyses
The amplified ITS1-5.8S rDNA-ITS2 region of the new Irpex species was 594
bp long. There was little intraspecific variation in the ITS region of I. lacteus
(mean value=0.40%, 0~3/436; 0~3 out of 436 positions showed nucleotide
difference) and in the ITS region of I. hydnoides (0.46%, 1~5/434). Average
sequence variation between I. hacksungii and I. lacteus or I. hydnoides was
3.05% (12~15/433) and 2.88% (12~13/434) respectively. These differences
Irpex hacksungii sp. nov. (Korea) ... 427
Irpex lacteus AF 163046 (Korea)
/, lacteus AF479666 (Korea)
lL. lacteus AY569563 (Canada)
‘| /. lacteus AY569564 (Canada)
|. lacteus AY569566 (Austria) [. lacteus
1. lacteus AY569565 (Germany)
I. lacteus ABO79264 (Japan)
e6/- /. lacteus ABO79265 (Japan)
|. lacteus AY569562 (Germany)
I. lacteus AY569561 (Austria)
=~ |/. lacteus ABO79266 (Japan)
lL. flacteus ABO79267 (Japan)
|. hydnoides AF479667 (Korea) I. hy dnoides
L hydnoides AF 479668 (Korea)
I. hacksungii SNU m-99032616 (Korea)
Ceriporia lacerata (EU301650)
—j1 change
FiGureE 2. The NJ tree of Irpex hacksungii, I. lacteus and I. hydnoides based on sequence data of
the ITS regions. Branches maintained in three different analyses (MP, ML and NJ analysis) were
presented by bold lines. Numbers above branches that are before the slash are NJ bootstrap
proportions and those that are after the slash are MP bootstrap proportions. Values below branches
are ML bootstrap proportions. The ITS sequence of Ceriporia lacerata (EU301650) was used as an
outgroup.
are greater than the variation observed between I. lacteus and I. hydnoides
(2.87%, 11~14/433). The aligned sequences of 15 Irpex species and Ceriporia
lacerata N. Maek., Suhara & R. Kondo (outgroup) formed a matrix with 604
nucleotide positions in length. Among these, 554 sites were constant, 38 sites
were variable but parsimony-uninformative, and 12 sites were parsimony-
informative. NJ, MP, and ML analyses yielded the same phylogenetic trees. The
_ NJ tree is presented in FiGurE 2. Ceriporia lacerata (DQ912694) was selected
as an outgroup, as justified in previous phylogenetic study (Lim & Jung 2003).
_ The tree showed that the new Irpex species was included in a monophyletic
_ clade with the two other Irpex species, I. lacteus and I. hydnoides. However, its
_ phylogenetic position was distinct within the Irpex clade (Fic. 2). Two Japanese
I. lacteus were grouped into I. hydnoides. They might be I. hydnoides because
i) I. lacteus had been treated as a monotypic genus for a long time and ii)
I. hydnoides was reported only recently (Lim & Jung 2003).
428 ... Lee, Kim & Lim
Discussion
Ryvarden (1991) classified the Polyporaceae into eleven groups and included
Irpex in the Junghuhnia group along with ten other related genera. However,
recent phylogenetic studies using rDNA sequence suggested that the genus
Irpex showed more relationship with the genera, Ceraceomyces Jiilich, Ceriporia
Donk, Ceriporiopsis Domanski, Cystidiodontia Hjortstam, Hexagonia Fr. and
Oxyporus (Bourdot & Galzin) Donk (Ko et al. 2001, Lee 2006). I. lacteus
and species from the genus Ceriporia have received greater attention as they
are important white rot fungi due to their production of endo- and exotype
cellulases (Hamada et al. 1999, Hoshino et al. 1994), antibiotics and irpexans
(Silberborth et al. 2000). They are also capable of decolorizing chemically
complex dyes and bioremediation of hazardous organic compounds (Kim &
Song 2000, Novotny et al. 2000).
Since Maas Geesteranus’s (1974) revision, two species have become accepted
in the genus Irpex: I. lacteus and I. hydnoides. We include one more species,
I. hacksungii, in this genus. Morphological characteristics and phylogenetic
analysis of the ITS region show that I. hacksungii is closer to I. hydnoides than to
I. lacteus. However, the new species is quite distinct from I. hydnoides, because
it possesses short teeth, and small lamprocystidia and basidiospores (TABLE 2).
In the molecular phylogenetic analysis, the three Irpex species formed a
monophyletic clade, but were separated into independent lineages. Additionally
I. lacteus is distributed worldwide as a cosmopolitan species whereas I. hydnoides
and I. hacksungii appear to be distributed in Far East Asia. I. hydnoides was
found in China recently (Dai et al. 2008) and Japan (Fie. 2). I. hydnoides and
I. hacksungii might be derived from I. lacteus and may have a potential for
biotechnological applications, similar to or higher that of I. lacteus.
Acknowledgments
This work was supported by the Korean indigenous species research project from the
National Institute of Biological Resources (NIBR), Ministry of Environment, Republic
of Korea. We are grateful to Dr. Rimvydas Vasaitis and Dr. Yu-Cheng Dai for pre-
submission review of the manuscript and Dr. Shaun Pennycook for improving the
English text.
Literature cited
Cejp K. 1931. Notes on Iowa Species of the Genus Irpex. Mycologia 23: 130-133.
Dai YC, Xiong HX. 2008. Irpex (Basidiomycota, Steccherinaceae) in China. Mycosystema 27:
515-519.
Gilbertson RL, Ryvarden L. 1986. North American Polypores. Volume 1. Fungiflora: Oslo
(Norway).
Irpex hacksungii sp. nov. (Korea) ... 429
Hamada N, Fuse N, Shimosaka M, Kodaira R, Amano Y, Kanda T, Okazaki M. 1999. Cloning and
characterization of a new exocellulase gene, cel3, in Irpex lacteus. FEMS Microbiology Letters
tye: 231-237.
Hoshino E, Kubota Y, Okazaki M, Nisizawa K, Kanda T. 1994. Hydrolysis of cotton cellulose by
exo- and endo-type cellulases from Irpex lacteus: differential scanning calorimetric study.
Journal of Biochemistry 115: 837-842.
Jeon YS, Chung H, Park S, Hur I, Lee JH, Chun J. 2005. jPHYDIT: a JAVA-based integrated
environment for molecular phylogeny of ribosomal RNA sequences. Bioinformatics 21:
3171-3173.
Kim HY, Song HG. 2000. Comparison of 2,4,6-trinitrotoluene degradation by seven strains of
white-rot fungi. Current Microbiology 41: 317-320.
Ko KS, Jung HS, Ryvarden L. 2001. Phylogenetic relationship of Hapalopilus and related genera
inferred from mitochondrial small subunit ribosomal DNA sequences. Mycologia 93:
270-276.
Largent DL, Johnson D, Watling R. 1977. How to identify mushrooms to genus, III. Microscopic
features. Mad River Press: Eureka (CA, USA).
Lee JS. 2006. Taxonomic and phylogenetic studies on Korean Aphyllophorales based on molecular
sequences. PhD Thesis. Seoul National University: Seoul (Korea).
Lee JS, Ko KS, Jung HS. 2000. Phylogenetic analysis of Xylaria based on nuclear ribosomal ITS1-
5.8S-ITS2 sequences. FEMS Microbiology Letters 187: 89-93.
Lim YW, Jung HS. 2003. Irpex hydnoides, sp. nov. is new to science, based on morphological,
cultural and molecular characters. Mycologia 95: 694-699.
Maas Geesteranus RA. 1974. Studies in the genera Irpex and Steccherinum. Persoonia 7: 443-581.
Novotny C, Erbanova P, Cajthamal T, Rothschild N, Dosoretz C, Sasek V. 2000. Irpex lacteus, a
white-rot fungus applicable to water and soil bioremediation. Applied Microbiology and
Biotechnology 54: 850-853.
Posada D, Crandall KA. 1998. MODELTEST: testing the model of DNA substitution. Bioinformatics
14: 817-818.
Ryvarden L. 1991. Genera of polypores: nomenclature and taxonomy. Fungiflora: Oslo (Norway).
Ryvarden L, Gilbertson RL. 1993. European Polypores, Part 1. Fungiflora: Oslo (Norway).
Silberborth S, Erkel G, Anke T, Sterner O. 2000. The irpexans, a new group of biologically active
metabolites produced by the basidiomycetes Irpex sp. 93028. The Journal of Antibiotics 53:
1137-1144.
Swofford DL. 2002. PAUP*4.0b Phylogenetic analysis using parsimony (* and other methods),
Version 4. Sinauer Associates: Sunderland (MA, USA).
White TJ, Bruns TD, Lee SB, Taylor JW. 1990. Amplification and direct sequencing of fungal
ribosomal RNA genes for phylogenetics. In PCR Protocols: A guide to methods and application
(eds. MA Innis, DH Gelfand, JJ Sninsky, TJ White), pp. 315-322. Academic Press: San Diego
HCA, USA).
MYCOTAXON
Volume 106, pp. 431-434 October-December 2008
Parmotrema sanctae-candidae,
a new species in the Parmeliaceae (lichenized Ascomycota)
from Brazil
SIONARA ELIASARO
sionara@ufpr.br
Depto. Botanica, Bioldgicas, Universidade Federal do Parana
Cx. P. 19031, 81531-970, Curitiba, PR - Brazil
Abstract — Parmotrema sanctae-candidae from Brazil is described as a new species in
the Parmeliaceae. It is similar to Parmotrema succinreticulatum but differs mainly in the
presence of marginal to submarginal isidia, which are often apically ciliate and become
granular and sorediate. It is so far only known from Parana state in southern Brazil.
Key words — lichens, Rimelia
Introduction
The genus Parmotrema A. Massal. includes species that have an upper cortex
consisting of a palisade plectenchyma or (rarely) paraplectenchyma with
vaults, have a pored or fenestrate epicortex, lack pseudocyphellae, may or may
not possess cilia, have laminal, perforate or eperforate apothecia, and usually
have simple rhizines and filiform, cylindrical, bacilliform or sublageniform
conidia (Blanco et al. 2005). There are 19 species of Parmotrema characterized
as having a maculate to reticulately cracked upper cortex and rhizines that
generally extend to the lobe margins. These species were previously included in
the genus Rimelia Hale & A. Fletcher.
During a study of the taxonomy and distribution of the Parmeliaceae in
Parana State, a new species of Parmotrema possessing a maculate to reticulately
cracked upper cortex was found. This species is described as new to science
based upon its morphological and chemical characteristics.
Materials and methods
The new species was described from specimens collected in the remnants
of an Atlantic mixed forest (Araucaria forest), a forest typical of southern
Brazil. The specimens were examined using standard stereoscopic and light
432 ... Eliasaro
microscopic procedures. Chemical constituents were identified by thin layer
chromatography (Culberson & Ammann 1979, Elix & Ernst-Russell 1993) and
by comparison with authentic samples.
Taxonomic description
Parmotrema sanctae-candidae Eliasaro, sp. nov. FIGURE 1
MycoBank MB 512333
Thallus chloroleucus, membranaceus, laxe adnatus, ad corticem arborum, usque 6 cm
diametri. Lobi irregulari, superpositi laterale, 3.0-8.0 mm latae, apex rotundus, margines
integrae, crenatae ad laciniatas, laciniae ascendentes, breves; cilia moderate densa,
nigra, praecipue simplex, 0.5-2.5 mm longitudinis. Superficies superna levis, notate
maculata reticulata, aliquando pruinosa alba in apicem loborum. Isidia marginalia ad
submarginalia in breves lacinias marginales, brevia, cilindrica, usque 0.5 mm altitudinis,
apex cataneus saepe ciliatus, convertus in granulatum sorediosum. Medulla alba.
Superficies infera nigra, polita, levis; margo castanea atra aut aliquando variata eboris
in lacinulas cum propaginibus, 1-2 mm diametri, levis, polita, rhizinata usque marginem
aut margo papillata; rhizinae nigrae, praecipue simplex ad-rare squarrosas, 0.1-3.0
mm, abundantes, similiter sparsae. Apothecia et pycnidia ignota. Atranorinam, acidum
fumarprotocetraricum et acidum succinprotocetraricum continens.
Type: BRAZIL. PARANA: General Carneiro. FAZENDA SANTA CANDIDA, 30.X.2004,
S Eliasaro 2798 (HOLOTYPE-UPCB).
ETyMOLoGy: From Sancta Candida, the place of origin of the type material.
Thallus greenish grey, membranaceous, loosely adnate, corticolous, up to
6 cm wide. Lobes irregularly branched, overlapping laterally, 3.0-8.0 mm
wide, apices rounded, margins entire, crenate to sublaciniate; cilia moderately
dense, black, simple, 0.5-2.5 mm long. Upper surface smooth, dull, strongly
reticulately maculate, sometimes pruinose towards the tips. Isidia marginal to
submarginal on short marginal laciniae, short-cylindrical, up to 0.5 mm long,
brown-tipped often apically ciliate, ultimately becoming granular and sorediate.
Medulla white. Lower surface black, glossy, smooth; marginal zone dark brown
or sometimes mottled ivory below the isidiate laciniae, 1-2 mm wide, rhizinate
to the margin or with a papillate zone along the margins; rhizines black, simple
to rarely squarrose, 0.1-3.0 mm long. Apothecia and pycnidia not seen.
CHEMISTRY: cortex: K+ yellow (atranorin); medulla: K+ faint yellow turning
brown, C-, KC+ faint yellow, P+ orange, UV- (succinprotocetraric and
fumarprotocetraric acids).
ADDITIONAL SPECIMENS EXAMINED — BRAZIL. PARANA: General Carneiro. FAZENDA
SANTA CANDIDA, 30.X.2004, S Eliasaro 2794 (UPCB).
Parmotrema sanctae-candidae is known only from the type locality, an Araucaria
forest in Parana State, southern Brazil.
COMMENTS — Parmotrema sanctae-candidae is distinguished by its distinct
reticulately maculate upper surface, its chemistry, and by its isidia. It is closely
Parmotrema sanctae-candidae sp. nov. (Brazil) ... 433
FiGuRE 1. Parmotrema sanctae-candidae (holotype in UPCB) Bar = 9 mm.
related to Parmotrema succinreticulatum (Eliasaro & Adler) O. Blanco et al.,
which also contains succinprotocetraric acid and fumarprotocetraric acid in
the medulla. The main taxonomic difference between the species is the presence
of isidia in Parmotrema sanctae-candidae.
The only species with which P sanctae-candidae might be confused is
P. subisidiosum (Mill. Arg.) Hale, which differs in containing salazinic acid
(medulla K+ yellow turning red) and in having cylindrical isidia that do not
become sorediate.
Acknowledgements
The author wishes to thank Prof. Nasser K. Hammad for checking the Latin descriptions,
Dr. John A Elix and Dr. Thomas Nash II] for the critical revision of the manuscript, and
“Instituto ECOPLAN” for collection permits.
434 ... Eliasaro
Literature cited
Blanco O, Crespo A, Divakar PK, Elix JA, Lumbsch HT. 2005. Molecular phylogeny of parmotremoid
lichens (Ascomycota, Parmeliaceae). Mycologia 97(1) 150-159.
Culberson CK Ammann K. 1979. Standarmethode zur Diinnschicht-chromatographie von
Flechtensubstanzen. Herzogia 5: 1-24.
Elix JA, Ernst-Russell KD. 1993. A catalogue of standardized thin layer chromatographic data and
biosynthetic relationships for lichen substances. 2" edition, Australian National University,
Canberra.
MY COTAXON
Volume 106, pp. 435-439 October-December 2008
Canoparmelia sanguinea,
a new Parmeliaceae from Brazil
MICHEL N. BENATTI & MARCELO P. MARCELLI
michel_benatti@yahoo.com.br
Instituto de Botanica, Secao de Micologia e Liquenologia
Caixa Postal 4005, Sao Paulo/SP, CEP 01061-970, Brazil
JOHN A. ELIx
John.Elix@anu.edu.au
Department of Chemistry, Building 33, Australian National University
Canberra, A.C.T. 0200, Australia
Abstract — Canoparmelia sanguinea, a Brazilian species of Parmeliaceae from Sao
Paulo State, is described as new to science.
Key words — Canoparmelia caroliniana, Canoparmelia concrescens, evernic acid
Introduction
Canoparmelia Elix & Hale, a segregate of the eciliate genus Pseudoparmelia
Lynge (Hale 1976), is characterized by typically gray or rarely yellow-green
thalli containing cortical atranorin and chloroatranorin [or rarely usnic acid],
3.0-5.0 mm wide, rotund or subrotund, eciliate lobes, a white medulla, a black
lower surface with naked brown margins and simple concolorous rhizines,
small ellipsoid ascospores 10-14 x 6-8 um, and fusiform or bifusiform conidia
7-10 um in length (Elix et al. 1986, Elix 1993).
The new species is based on specimens collected in the State Forest Reserve
of Serra da Cantareira (23° 20’—23° 27’S, 46° 28’'-46° 42’W), in an open area
near the Atlantic Rainforest in the very north of Sao Paulo city, and adjacent
municipalities.
Material and methods
The morphological characters were examined under a stereo-microscope.
Anatomical sections were made with razor blades. The chemical constituents
436 ... Benatti, Marcelli & Elix
were analyzed by color reaction spot tests including potassium hydroxide (K),
sodium hypochlorite (C) and paraphenylenediamine (P). All specimens were
tested under UV light, and chemical constituents were identified by thin-layer
chromatography (TLC) using solvent C (Bungartz 2001), high performance
liquid chromatography (HPLC) (Elix et al. 2003) and comparison with
authentic samples.
In the description, the Latin diagnosis refers exclusively to the holotype and
the English text includes data from all the specimens examined.
Canoparmelia sanguinea Marcelli, Benatti & Elix, sp. nov. FIGURE 1
MB 511928
DiaGnosis: Species thallo similis Canoparmelia caroliniana sed obscurior, superus
rugosus vel scrobiculatus, distincte maculatus, isidiis fuscatis vel apicibus brunneis,
simplicibus vel parum ramosis, arcte aggregatis, subtus niger vel atrobrunneus, medulla
C+ sanguineus, KC+ sanguineus et acidis olivetolcarboxylico, 4-O-methylolivetolcarbox
ylico, decarboxyanziaico, decarboxystenosporico, decarboxyperlatolico, divaricatinico et
glomelliferico, glomellino, et depsidellino B continens differt.
Ho.Lotype—Brazil, Sao Paulo State, Municipality of Mairipora, Cantareira Range, on a
tree trunk felled during village construction, leg. M.P. Marcelli 6029, 03-XI-1989 (SP).
THa.LLus sublobed, dark cinereous green becoming dusky green in the
herbarium, up to 9.0 cm diam., submembranaceous, corticolous. Lobes
irregularly ramified to subdichotomous, 1.0-3.5 mm wide, contiguous to
imbricate in the younger parts and becoming somewhat crowded in the center,
closely adnate, with subrotund or sometimes subtruncate apices, the margins
entire to partly subcrenate, frequently involute, the upper surface continuous
to irregularly cracked, often rugose or slightly scrobiculate especially in
the older parts, tsmooth in the distal parts. MACULAE irregularly linear,
laminal, becoming confluent and more conspicuous near the lobes apices,
sometimes forming inconspicuous fissures. CILIA absent. LACINULAE absent,
but sometimes forming small, simple, flat, subtruncate adventitious lobules in
the older parts, 0.3-1.2 x 0.3-1.6 mm. SOREDIA and PUSTULES absent. IsIDIA
laminal, granular to short cylindrical, 0.05-0.50 x 0.05-0.10 mm, smooth,
simple to sparsely branched, generally densely clustered, not coralloid; erect,
smooth, firm, partially darkened or with brown apices, eciliate. MEDULLA
white. LOWER SURFACE mottled, black to dark brown, shiny to opaque, smooth
to rugose, sometimes slightly veined or papillate. MARGINAL ZONE brown,
shiny, 0.5-1.5 (—4.0 at the lobes apex) mm wide, less evident in some parts,
smooth to rugose or veined, usually naked but sometimes sparsely papillate or
rhizinate approaching the center. RHIzINAE dark brown to black, but brown or
white near the marginal zone, simple to rarely furcate, acute, (0.10—) 0.20-0.70
(-1.00) x ca. 0.05 mm, frequent, tevenly distributed. APOTHECIA not seen.
PYCNIDIA rare, submarginal, with black ostioles. CONIDIA not seen.
Canoparmelia sanguinea, sp. nov. (Brazil) ... 437
FiGure 1. The holotype (a single specimen) of Canoparmelia sanguinea (2),
Bar = 1 cm.
438 ... Benatti, Marcelli & Elix
SPOT TESTS: cortex K+ yellow, UV-; medulla K—-, C+ reddish rose, KC+ red, P+
(?) very faint yellowish, UV-.
TLC/HPLC: cortical atranorin (minor); medullary olivetolcarboxylic acid
(minor), 4-O-methylolivetolcarboxylic acid (minor), glomelliferic acid
(minor/absent), glomellin (trace/minor), decarboxyanziaic acid (major),
decarboxystenosporic acid (minor), decarboxyperlatolic acid (minor),
divaricatinic acid (minor), depsidellin B (minor), unknowns (minor).
PARATYPE-Brazil, Sao Paulo State, Municipality of Mairipora, Cantareira Range, on a
tree trunk brought down during village building, leg. M.P. Marcelli 6028, 03-XI-1989
(B, SP).
COMMENTS— Canoparmelia sanguinea is an isidiate species which has a dark
gray green thallus, imbricate or crowded lobes, often with involute margins, an
almost entirely subrugose or slightly scrobiculate upper surface with prominent
maculae, especially near the lobes tips, and a mottled dark brown to black lower
surface. The rhizines are commonly paler in distal areas.
Due its size, overall thallus shape, the presence of isidia and prominent
macules, in the field C. sanguinea could be mistaken for darkened specimens of
the very common C. caroliniana (Nyl.) Elix & Hale.
However, its medullary chemistry is unique and very different from
C. caroliniana, as was first noted by the immediate C+ and KC+ red spot tests
[which in Parmeliaceae usually indicate the presence of the lecanoric/gyrophoric
acids, olivetoric acid or anziaic acid complexes] instead of the C- and KC+ faint
violet/rose reaction of C. caroliniana (due to the presence of perlatolic acid).
The new species produces dense, intensely clustered and ramified isidia
with brown apices (which give the false impression of being coralloid), so
that the isidia appear darker and more dense by comparison with those of
C. caroliniana.
According to Hale (1976) C. concrescens (Vain.) Elix & Hale, another similar
species, differs in having a flat, continuous, emaculate upper surface and in
producing only divaricatic acid in the medulla.
This species is named after the red C and KC spot tests.
Acknowledgments
The authors wish to thank Harrie J. M. Sipman (Berlin) and Richard Harris (New York)
for the critical revision of the manuscript and help with the Latin diagnosis.
This work could not have been accomplished without the support of the FAPESP
(Fundacao para o Amparo a Pesquisa do Estado de Sao Paulo) for Scientific Initiation
grant (n° 00/01009-1) and the Conselho Nacional de Desenvolvimento Cientifico e
Tecnoldgico (CNPq) — for the research support to the second author.
Canoparmelia sanguinea, sp. nov. (Brazil) ... 439
Literature cited
Bungartz F. 2001. Analysis of lichen substances. In http://nhc.asu.edu/lichens/lichen_info/tle.
jsp#TLC2. Accessed on July 2008.
Elix JA. 1993. Progress in the generic delimitation of Parmelia sensu lato lichens (Ascomycotina:
Parmeliaceae) and a synoptic key to the Parmeliaceae. The Bryologist 96(3): 359-383.
Elix JA, Johnston J, Verdon D. 1986. Canoparmelia, Paraparmelia and Relicinopsis, three new genera
in the Parmeliaceae (Lichenized Ascomycotina). Mycotaxon 27: 271-282.
Elix JA, Giralt M, Wardlaw JH. 2003. New chloro-depsides from the lichen Dimelaena radiata.
Bibliotheca Lichenologica 86: 1-7.
Hale ME. 1976. A Monograph of the Lichen Genus Pseudoparmelia Lynge (Parmeliaceae).
Smithsonian Contributions to Botany 31: 1-62.
’
MYCOTAXON
Volume 106, pp. 441-444 October-December 2008
Two new species of Pertusaria (Pertusariaceae) from China
QIANG REN, ZHONG-SHUAI SUN & ZUN-TIAN ZHAO*
rengiang@sdnu.edu.cn sun2143998@yahoo.com.cn *ztzhao@sohu.com
College of Life Sciences, Shandong Normal University
Jinan, 250014, China
Abstract -- The species Pertusaria qilianensis and Pertusaria paragilianensis from
Northwestern China are described as new to science.
Keywords —- lichen, taxonomy, planaic acid
Introduction
The lichen genus Pertusaria in China has received little systematic study. All
the taxa reported before 2004 have been revised, and the number of accepted
taxa reduced to 45 (Zhao et al. 2004). A study of several recent collections of the
genus from northwestern China has revealed the presence of two new species
which are described here.
Material and methods
The specimens examined were collected in Qilian Mountains and Mt. Helan,
Northwestern China, and are preserved in SDNU (the Lichen Section of
Botanical Herbarium, Shandong Normal University).
A dissecting microscope (Motic K-400L) anda research microscope (JNOEC
XS-213) were used for the morphological and anatomical studies. The lichen
substances was detected by thin-layer chromatography (TLC) (Elix & Ernst-
Russell 1993) and high performance liquid chromatography (HPLC) (Elix et
al. 2003).
Taxonomic description
Pertusaria qilianensis Q. Ren & Z.T. Zhao, sp. nov. FIGURES 1-2
MycoBankK MB 512348
Thallua crustaceus, cineraceus vel flavo-cineraceus, tenuis, subrimosus, superficies
scaber et rugosus, isidiis et sorediis destitutis, corticola. Apothecia disciformia, numerosa
* Corresponding author
442 ... Ren, Sun & Zhao
et dispersa, 0.7-1.5 mm diam., disci nigri et sorediati, marginibus thallo concolores,
laceratus et lobularis. Asci clavati, ascosporae 2nae, ellipsoideae ad ovoideae, (85-)100-
180 um longae, (40-)50-80 um latae, parietibus laevis, 7-13 um crassis. Thallus acidum
planaicum continens.
Typus: CHINA. Qinghai provincia, Qilian, Niuxinshan, alt. 3120m, ad corticem arborum,
H. Y. Wang 20071461-1. (holotypus in SDNU).
ETyMo.ocy: The specific epithet gilianensis is derived from the type locality in Qilian
Mountains.
THALLUS crustose, thin, very pale gray to yellowish gray, slightly cracked, surface
matt and rugose; lacking isidia; corticolous; APOTHECIA disciform, numerous,
scattered, sometimes confluent, sub-immersed to sessile, constricted at the
base, 0.7-1.5 mm diam., discs black, sorediate, margins lacerated and stellate,
concolorous with the thallus; asci cylindric, 210-315 x 50-100 um, 2-spored;
ASCOSPORES ellipsoid to ovoid, hyaline, (85—)100-180 um long, (40-)50-80
uum wide, double-walled, smooth, 7-13 um thick; pyCNIDIA not seen.
CHEMISTRY: K-, C-, KC-, P-, UV-; planaic acid (TLC and HPLC).
DISTRIBUTION AND SUBSTRATE —Pertusaria qilianensis is a corticolous species,
found in northwestern China at elevations of 2670 to 3500 m.
SPECIMENS EXAMINED — CHINA. QINGHAI: Qilian Co. BINGGOU- XIAXIGOU, alt.
3500m, on bark 10/viii/2007, X. L. Shi 20071426, 20071429. NIUXINSHAN, alt. 3050m,
on bark 11/viii/2007, H. Y. Wang & X. L. Lu 20071462, 1470-1; H. Y. Wang 20071461-1,
20071465-1; Y. D. Du 20071448-1; alt. 3100m, on bark 11/viii/2007, H. Y. Wang 20071485-
1, 20071496; Y. D. Du 20071520-3; Z. S. Sun 20071487-1, 20071504-1; alt. 3150m, on bark
11/viii/2007, Z. S. Sun 20071534-2; alt. 3200m, on bark 11/viii/2007, Y. D. Du 20071574-
2; J. G. Liu 20071534-2; alt. 3250m, on bark 11/viii/2007, Z. S. Sun 20071583-1; X.L. Shi
20071587-1; alt. 3300m, on bark 11/viii/2007, Y. D. Du 20071594;alt. 3400m, on bark
11/viii/2007, X. L. Shi 20071620. NinextA: Mt. Helan suyUKOU-BEIGOU, alt. 2670m, on
bark 22/viii/2007, E Yang 20072545; (SDNU).
COMMENTS — Pertusaria gilianensis is characterised by asci with two smooth
ascospores, sorediate disciform apothecia and the presence of planaic acid.
Planaic acid is an uncommon depside in the genus Pertusaria and usually
present as only a trace compound. ‘The only species in which planaic acid was
reported as a major component were P. leucothelia, P. planaica and P. siamensis.
The new species can be readily distinguished from the former two species by
the number of ascospores. In the new species there are two ascospores per
ascus in contrast to the four-spored asci in PB leucothelia and the eight-spored
asci in P. planaica. The new species differs from P. siamensis in morphology,
especially the development of apothecia, disciform apothecia compared to
conspicuous verruciform apothecia in P siamensis (Archer & Elix 1992, Archer
1997, Jariangprasert 2005).
Pertusaria spp. nov. (China) ... 443
Figures 1-2. P. gilianensis. 1. Morphology (bar = 1mm); 2. Apothecium cross section
(bar = 100 um). Ficures 3-4. P. paragilianensis. 1. Morphology (bar = 1mm); 2. Apothecium
cross section (bar = 100 tm).
_ Pertusaria paragilianensis Z.S. Sun & Z.T. Zhao, sp. nov. FIGURES 3-4
MycoBAnk MB 512347
Thallua crustaceus, cineraceus, crassiusculuss, surareolatus, superficies laevis et nitida,
isidiis et sorediis destitutis, corticola. Apothecia disciformia, numerosa et dispersa, 0.5-
1.2 mm diam., disci nigri, marginibus thallo concolores, crassus et inflated. Asci clavati,
ascosporae 2nae, ellipsoideae ad ovoideae, 90-130 um longae, 45-70 um latae, parietibus
laevis, 7-9 um crassis. Thallus acidum planaicum continens.
Typus: CHINA. Qinghai provincia, Qilian, Niuxinshan, alt. 3120m, ad corticem arborum,
Z. S. Sun 20072496. (holotypus in SDNU).
Erymo_oey: from the Greek para, near, a reference to the similarity of the new species
to Pertusaria qilianensis.
THALLUS crustose, pale gray to gray, moderately thick, slightly tuberculate,
surface smooth and shiny; lacking isidia and soredia, corticolous; APOTHECIA
disciform, conspicuous, numerous, scattered, sometimes confluent, sub-
- immersed to sessile, 0.5-1.2 mm diam., concolorous with the thallus, discs
black, epruinose, margins thick and irregular inflated; asci cylindric, 200-235
x 50-85 um, 2-spored; ascosporss ellipsoid to ovoid, hyaline, 90-130 um long,
45-70 um wide, double-walled, smooth, 7-9 um thick; PYCNIDIA not seen.
444 ... Ren, Sun & Zhao
CHEMISTRY: K-, C-, KC-, P-, UV-; planaic acid (TLC and HPLC).
DISTRIBUTION AND SUBSTRATE —At present, Pertusaria paragilianensis is
known from Qilian mountains, northwestern China at elevations of 3120 to
3200 m; on bark.
SPECIMENS EXAMINED — CHINA. QINGHAI: Qilian Co. NIUXINSHAN, alt. 3200m,
on bark 11/viii/2007, J. G. Liu 20071572-4; alt. 3120m, on bark 12/viii/2007, Z. S. Sun
20072496 (SDNU).
COMMENTS — Pertusaria paragilianensis is characterised by asci with two
smooth ascospores, inflated apothecia margin and the presence of planaic
acid. The new species is chemically identical to P ginghaiensis but that species
has a conspicuous rough thallus surface, sorediate apothecia and a lacerated
apothecia margin. The new species resembles the North America species
P. saximontana but differs from that species by chemistry and the apothecia
margin. P saximontana has crenulate or eroded apothecium margins and
contains 2-O-methylconfluentic acid as its major compound, whereas the new
species has inflated apothecia margins and contains planaic acid (Dibben 1980,
Lumbsch et al. 1999).
Acknowledgements
The authors are grateful to Professor J.A. Elix, (Australian National University, Canberra)
for the liquid chromatography results. The project was financially supported by the
National Natural Science Foundation (30870012) and Shandong Provincial Natural
Science Foundation (Y2007D21) of China. The authors thank Dr. Alan W. Archer and
Dr. André Aptroot for expert presubmission review.
Literature cited
Archer AW. 1997. The lichen genus Pertusaria in Australia. Biblioth. Lichenol. 69: 1-249.
Archer AW, Elix JA. 1992. Further new species and new reports of Pertusaria (lichenised
Ascomycotina) from Australia. Mycotaxon 43: 417-431.
Dibben MJ. 1980. The chemosystematics of the lichen genus Pertusaria in North America north of
Mexico. Milwaukee Publi. in Biol. Geol. 5: 1-162
Elix JA, Ernst-Russell KD. 1993. A catalogue of standardised thin layer chromatographic data
and biosynthetic relationships for lichen substance, 2" Edition. 163 pp. Australian National
University, Canberra.
Elix JA, Giralt M, Wardlaw JH. 2003. New chloro-depsides from the lichen Dimelaena radiata.
Biblioth. Lichenol. 86: 1-7.
Lumbsch HT, Nash TH IH, Messuti MI. 1999. A revision of Pertusaria species with hyaline
ascospores in southwestern North America (Pertusariales, Ascomycotina). The Bryologist 102:
2159-239.
Jariangprasert S, Anusarnsunthorn V. 2005. Additional new taxa in the lichen genus Pertusaria
(lichenized Ascomycota) from Thailand. Mycotaxon 91: 279-292.
Zhao ZT, Ren Q, Aptroot A. 2004. An annotated key to the lichen genus Pertusaria in China.
Bryologist 107: 531-541.
MYCOTAXON
Volume 106, pp. 445-448 October-December 2008
The new lichen species Endocarpon crystallinum
from semiarid deserts in China
JUN YANG & JIANG-CHUN WEI*
weljc2004@126.com
Key Laboratory of Systematic Mycology e& Lichenology
Institute of Microbiology, Chinese Academy of Sciences
No. A3 Datun Road, Chao-yang District, Beijing 100101, P.R.China
Abstract — The newlichen species Endocarpon crystallinum was collected from semiarid
deserts in China and is described in the present paper. Research on the morphology,
anatomy, and chemistry of the species has been carried out. The species is unique in
the genus by the thick upper cortex that is broken into columnar polygonal elements.
A similar cortex structure is however known from a few, unrelated, other desert lichens
and appears to be an adaptation to that climate.
Keywords — Ascomycota, cortex structure, Verrucariaceae
Introduction
During research on lichen species and their genetic diversity in the arid and
semiarid deserts of northwestern China, numerous species of Endocarpon
have been found. The genus is mainly characterized by a squamulose thallus,
immersed perithecioid ascomata with hymenial algae, and asci with (1-)2(-8)
muriform ascospores. Among the specimens collected was one that could not
be identified with any of the known species, and it is consequently described
below as new to science. The genus Endocarpon belongs to the Verrucariaceae
(Kirk et al. 2001), Verrucariales, Chaetothyriomycetidae, Eurotiomycetes,
Ascomycota (Geiser et al. 2006).
Materials and methods
The specimens examined were collected by either J. Yang and E.R. Zhang or
T. Zhang from the deserts in the Shanxi province in northwestern China.
Hand sections and squash preparations were made for anatomical study.
The morphological and anatomical examinations were performed using the
* Corresponding author.
446 ... Yang & Wei
dissecting microscope Leica MZ8 and the biological microscope Leica DMR
respectively. For chemical analysis, the standardized thin-layer chromatograph
(TLC) method for identification of lichen products was used, applying solvent
C (Culberson 1972, Culberson & Kristensson 1970, White &James 1985).
Results and discussion
Endocarpon crystallinum J.C.Wei & Jun Yang, sp. nov. Fic. 1
MycoBank MB512364
Species nova cortice supero crasso quo fracto in elementa similia crystallis, sed constantia
ex textura paraplectenchymata hyalina a congeneribus diversa.
Typus: China, Shanxi provincia: Yang-gao comitatus, Xiejiatun pagus, ad arenam,
September 23, 2004, J. Yang & E.R. Zhang SX-28 (holotypus) in HMAS-L conservatur.
EtyMoLoGcy: the specific epithet “crystallinum” refers to the upper cortex of the thallus,
which is covered with a broken hyaline layer reminiscent of crystals.
EXPANDED DESCRIPTION — ‘thallus terricolous, squamulose, consisting of
small squamae, covered with a thick layer of hyaline tissue that is broken into
columnar polygonal elements resembling crystals (Fic. 1 A); squamae solitary,
with dark brown to black peripheral margins (Fic. 1 A), or occasionally
contiguous, never imbricate, pale to yellowish brownish, rounded, elongate or
irregular, shallowly to deeply lobate, 2-4(-7) mm broad.
Upper cortex consisting of two layers: the upper thick paraplectenchymatous
layer of hyaline tissue that is broken into columnar polygonal elements
resembling crystals in the shape of peaks is (30-)50-175 um tall, the elements
(50-)62.5-275(-475) um wide at the base, 17.5-175(-225) um wide at the
top; the lower brownish paraplectenchymatous layer is 34-80(-110) um tall.
(Fic. 1 B)
Algal layer 38-100(-112)um tall, consisting of subglobose and bright green
cells, 5—7.5 x 5-10 um, without visible medulla (Fic. 1 B).
Lower cortex consisting of obscure plectenchyma, dark brown to black,
about 50 um tall, with brown to black and branched rhizines of 1.5-2.5 mm
long (Fre. 1 C).
Ascomata perithecioid, subglobose, uniloculate, immersed in thallus,
solitary, 1-6 per squama, with brown to dark brown apex of 0.25 mm in diam.
bearing a depressed ostiole lighter in color than the surrounding tissue; centrum
obpyriform, 150-490 um wide; excipulum brown to dark brown, 105-400 um
thick at the base and sides, pale brown at the apex near the ostiole; gelatinizing
periphysoids 75-100 um long, simple to sparingly branched; hymenial gel
Lugol's I + blue; hymenial algal cells globose to oval, green, 2.5-5 um in diam.
or 2,5—-3.5 x 3.5—-5 um (Fics. 1D, E).
Asci fissitunicate, bisporous (Fic. 1E), clavate to cylindro-clavate, 40-60 x
15.5-21.5 um; ascospores muriform, yellowish to brownish, elongate-ellipsoid
|
|
:
Endocarpon crystallinum sp. nov. (China) ... 447
BUEN
a
Fig. 1. A. Endocarpon crystallinum: Upper surface of squamae covered with minute crystal-like
broken cortex elements and a few perithecioid ascomata of which only the ostioles are visible (bar
= 1mm); B. Cross section of a squama (bar = 50u1m); C. Cross section of a squama with branched
thizines (bar=l1mm); D. The hymenial algal cells and ascospores within a perithecioid ascoma
(bar = 100um); E. An ascus containing 2 spores within a perithecioid ascoma; note that the upper
spore is much broader and shorter than the lower spore (bar = 20 um); F. A muriform ascospore
(bar =20um).
(a = algal layer; as = ascus; c = crystals; ha = hymenial algal cells; o = ostiole; psd = periphysoid;
r= rhizines; s=spore; t=thallus)
to subcylindrical, with 2-4 transverse divisions and 9-15 longitudinal divisions,
the upper spore much broader and shorter than the lower spore; upper spore:
50-65(-87.5) x 12.5-15um; lower spore: (32.5-)40-50 x15-17.5um (Fics. 1
PF).
Conidiomata not seen.
CHEMISTRY: no lichen substances detected with TLC.
448 ... Yang & Wei
ADDITIONAL SPECIMENS EXAMINED: China, Shanxi prov.: Yang-gao county: Xiejiatun,
on the wall of soil, September 23, 2004, J. Yang & E.R. Zhang SX-32 (HMAS-L); west of
the Yang-gao county town 2 km, on slope of soil, April 20, 2007, J. Yang & T. Zhang,
SX-093, SX-113 (HMAS-L); Ningxia Autonomous region, Xiang mountain of Shapotou
region, soil among rocks; April 17, 2007, J. Yang & T. Zhang SPT-202 (HMAS-L).
CoMMENTs: This new species resembles in its upper cortex structure several
other unrelated lichens from desert habitats. Examples are Psora crystallifera
and several Toninia species (Timdal 1991), e.g. T. albilabra and T. sculpturata.
The congruence is so striking that Fig. 2E in Timdal (1991), taken from Toninia
albilabra, is indistinguishable from Fic. 1B below, taken from the new species,
which belongs to a different order (and even class). This structure is undoubtedly
an adaptation to desert climate.
The new species differs from other members currently accepted in the genus
(Breuss 2002, Harada 1993, McCarthy 1991) by the thick upper cortex that
is broken into columnar polygonal elements ses nang crystals, but made of
hyaline paraplectenchymatous tissue.
Acknowledgments
This work was financially supported by the National Natural Science Foundation of
China (NSFC, No.30270005) and the Ministry of Science and Technology of the People’s
Republic of China (MOST, No.2007AA021405). The authors are most grateful to Dr. A.
Aptroot and Dr. H. Sipman for reading the manuscript and giving the valuable comments
and suggestions, and also to Mr. E.R. Zhang and Mr. T. Zhang for accompanying with
the first author to do the field work.
Literature cited
Breuss O. 2002. Endocarpon. pp. 181-187 in: Nash TH, Ryan BD, Gries C & Bungartz E. Lichen flora
of the greater Sonoran desert region. Volume 1. Arizona State University, Tempe.
Culberson CFE. 1972. Improved conditions and new data for the identification of lichen products by
a standardized thin-layer chromatographic method. J. Chromatogr. 72:113-125.
Culberson CF, Kristensson H. 1970. A standardized method for the identification of lichen
products. J. Chromatogr. 46:85-93.
Geiser DM, Gueidan C, Miadlikowska J, et al. 2006. Eurotiomycetes: Eurotiomycetidae and
Chaetothyriomycetidae. Mycologia 98(6):1053-1064.
Harada H. 1993. A taxonomic study of the lichen genus Endocarpon (Verrucariaceae) in Japan.
Nova Hedwigia 45:335-353.
Kirk PM, Cannon PF, David JC, Stalpers JA. 2001. Ainsworth & Bisby’s Dictionary of the Fungi
(9th Edition) CABI Publishing, Egham.
McCarthy PM. 1991. The lichen genus Endocarpon Hedwig in Australia —- Lichenologist 23(1):
2/=52.
Timdal E. 1991. A monograph of the genus Toninia (Lecideaceae, Ascomycetes). Opera Botanica
110: 1-137,
White FJ, James PW. 1985. A new guide to microchemical techniques for the identification of lichen
substances. Brit. Lich. Soc. Bull. 57 (Suppl.): 1-41
MY COTAXON
Volume 106, pp. 449-453 October-December 2008
Gloeocantharellus persicinus,
a new species from China
CHUN- YING DENG & Tat-Hut Li*
mycolab@263.net
Guangdong Provincial Key Laboratory of Microbial Culture Collection & Application,
Guangdong Institute of Microbiology
Guangzhou 510070, China
Abstract — Gloeocantharellus persicinus is described as a new species from China
supported by macro- and microscopic photographs. It is compared with G. echinosporus,
G. lateritius, G. okapaensis, G. purpurascens and G. uitotanus. The holotype (GDGM
21480) is deposited in the Herbarium of Microbiology Institute of Guangdong Province
(GDGM).
Key words — taxonomy, Gomphaceae, cantharelloid fungus
Introduction
Gloeocantharellus Singer is a small genus of Gomphaceae with only 6 species
recognized (Singer 1945, Corner 1969, Petersen 1971, Vasco-Palcacios &
Franco-Molano 2005). The genus is characterized by its numerous gloeoplerous
hyphae, gloeocystidia, ornamented spores, and bifurcate lamellate to slightly
wrinkled hymenophore (Singer 1945, Corner 1969, Ainsworth et al. 1973). All
species of Gloeocantharellus are terrestrial and generally found associated with
ectomycorrhizal trees. Five species of Gloeocantharellus have been reported
from Asia, i.e. G. corneri (Singer) Corner, G. echinosporus Corner, G. lateritius
(Petch) Corner, G. okapaensis (Corner) Corner, and G. purpurascens (Hesler)
Singer (Lloyd 1917, Corner 1969, Pegler 1986, Yuan & Sun 1994, 2007).
However, the only Gloeocantharellus recorded for China — G. purpurascens
— should be excluded, because the field photographs (Yuan & Sun 1994,
2007) and re-examination of the specimens (HKAS 15708) showed that the
| basidiomata represented Gomphus clavatus (Pers.) Gray, not Gloeocantharellus
| purpurascens; furthermore, the cited specimens, with basidiospores 13-16 x
6-7.5 um, do not have any gloeocystidia.
*Corresponding author
450 ... Deng & Li
During a survey on the mycoflora of E-huangzhang Nature Reserve in Yangchun
County of Guangdong Province, a species belonging to Gloeocantharellus was
discovered and it possesses some distinguishing features from any known
species. A new species is therefore proposed as follows.
Materials and metheds
Specimens were annotated and photographed in the field, dried in an electric
drier, and then deposited in herbarium. Tissues were mounted in 5% KOH for
microscopic examination. The lengths and widths of 20 spores were randomly
selected, and Q represents the mean length/width of the 20 spores. The
microscopic characters were observed under light microscope and scanning
electronic microscope. Colour references are presented according to Kornerup
& Wanscher (1978). The holotype has been deposited in the Herbarium of
Guangdong Institute of Microbiology (GDGM).
Taxonomy
Gloeocantharellus persicinus T.H. Li, Chun Y. Deng & L.M. Wu, sp. nov.
MycoBank MB 511995 Figs. 1-5
BASIDIOMATA 4-6 cm alta, gomphoidea vel cantharelloidea cum hymenophora lamellata.
PrLEus 3.8-7.0 cm x 3.5-6.0 cm, convexus dein applanatus vel infundibuliformis,
viscidulus, dilute aurantiacus, rufulo-aurantiacus vel persicinus; margine undulatus
vel valvatus, primo incurviusculus. CONTEXTUS albus, tactu immutabilis. LAMELLAE
albidae, flavidae vel cerinoflavidae cum coloratione olivaceo-grisea, subconfertae, 9-12
perm, < 3.5 mm latae, decurentes, inaequales, saepe furcatae. STIPES 4-4.5 cm x 0.9-1.1
cm, cylindraceus, basim versus attenuatus, centralis vel eccentricus, concolor cum pileo
vel pallidulus quam pileus. BASIDIOSPORAE (7.8-)8.6-10.1 x 4.1-5.8 um, ellipsoideae,
verruculosae, ochraceae, inamyloideae. BAsiDIA 36-50(-70) x 6-10 ym, clavata,
2-4-sterigmata 5-8 um longis. GLoEocysTIDIA lanceolata vel subclavata, hymenialia,
42-80(-160) x 4.5-9 um. PILEIPELLIS et STIPITIPELLIS ixocutis, cum hyphis oleiferis.
HYPHAE OLEIFERAE 2.9-7.3 um latae. HYPHAE 2.9-5yum latae, fibulatae.
TYPE COLLECTION — CHINA, Guangdong, Yangchun County, 15 May 2004, T.H. Li &
L.M. Wu [GDGM 21480, HOLOTYPE].
BASIDIOMATA 4-6 cm tall, gomphoid to cantharelloid in general contour but
with obvious lamellate hymenophore. PILES 3.8—7.0 cm x 3.5-6.0 cm, convex
then plane to infundibuliform, usually depressed at centre, subviscid, light
orange or reddish orange to peach-colour (6A4, 7A4, 7A6, 8A4); margin wavy
or valvate, initially inflexed. CONTEXT white, bruising unchanged. LAMELLAE
whitish to pale yellow (4A1 to 4A3) tinted with olive-grey (4D2) , becoming
olive-grey, olive-brown to light brown (4D2, 4D4 to 5D4) when dried, crowded,
9-12 per cm at pileus-margin, <3.5 mm wide, deeply decurrent, unequal,
sometimes bifurcate but not or hardly anastomosing, edges even. STIPE 4-4.5
cm long, 0.9-1.1 cm thick at apex, cylindrical, equal to slightly enlarged
Gloeocantharellus persicinus sp. nov. (China) ... 451
AccoV SpotMagn Def WD
WOKV4G 6000x%. SE 9.2
2
Fics. 1-5: Gloeocantharellus persicinus (holotype). 1. Basidiomata; 2-3. Basidiospores
4. Basidia in hymenium; 5.Gloeocystidia in hymenium.
Bars: 1 = 1 cm; 2 = 5um; 3 = 5 um; 4= 10 um. 5 = 10 um.
downwards, tapered at base, central to slightly eccentric, mostly concolorous
with the pileus, peach-colour, pastel red to pale red (7A4,7A3), in some parts
paler or whitish.
BASIDIOSPORES (7.8—)8.6-10.1 x 4.1-5.8 um (x=9.7 x 5.2, n= 20, Q=1.86),
elongate ellipsoid, ornamented with strongly cyanophilic warts, yellow to
452 ... Deng & Li
cinnamon brown in mass, pale yellow to pale cinnamon in KOH under
microscope, inamyloid. Basip1A 36-50(-70) x 6-10 um, clavate, hyaline, with
2-4 sterigmata 5-8 um long. PILEIPELLIS an ixocutis of a compact layer of
repent and interwoven gelatinous hyphae with gloeoplerous hyphae. PILEus
CONTEXT of thin-walled, parallel to closely interwoven hyphae; gloeoplerous
hyphae frequent, tortuous, rarely branched, some terminating among basidia.
STIPE CONTEXT of packed hyphae of two types: (1) thin-walled, parallel, hyaline,
2.9-5 um in diam., and (2) gloeoplerous, tortuous, aseptate, interwoven, yellow,
2.9-7.3 um in diam., common and abundant. GLoEocysTIDIA 42-80(-160) x
4.5-9 um, completely immersed to slightly projecting beyond the hymenium,
thin-walled, lanceolate to subclavate, with yellow contents in H,O and KOH.
STIPITIPELLIS formed of parallel and narrow hyphae, with some gloeoplerous
hyphae. CauLocystTIpIA not observed. Clamp connections abundant.
HABITAT, DISTRIBUTION AND SEASON — Single to scattered, on soil in broad-
leaved forest; China (Guangdong). May.
COMMENTS—G. persicinus is similar in colour to three other pinkish to
orange red Asian species, i.e. G. echinosporus, G. lateritius, and G. okapaensis.:
However, the basidiospores of G. echinosporus are obviously spiny and wider;
in G. lateritius the basidiospores are larger (10-13.5 x 6-7 um) and the context
turns purplish on exposure; and in G. okapaensis the pileus is dry, the context
becomes dingy vinaceous mauve on bruising, and the basidiospores are also
larger (9-12 x 5-7 um) (Corner 1969, Pegler 1986). Macroscopically, the new
species resembles G. purpurascens, but that North American species has larger
basidiomata and distinctive purple or violet discoloration on the bruised parts
of the basidiome (Smith & Hesler 1943). Another red species, G. uitotanus
Vasco-Pal & Franco-Mol. originally described from Central America, can also
be distinguished, the pileus being deeper red to reddish brown (8C8-9C8),
the basidiospores larger [(8—)8.8-12 x (4-)4.8-6(-6.4) um], and the lamellae
narrower (up to 2 mm broad), fewer (subdistant), subdecurrent rather than
decurrent, and more strongly anastomosing than those of the new species
(Vasco-Palacios & Franco-Molano 2005).
Acknowledgements
The study was supported by the National Natural Science Foundation of China
(No. 30770004, 30870019) and the Natural Science Foundation of Guangdong (No.
E05202480). The authors are very grateful to Dr. Roy Watling of Royal Botanic Garden
Edinburgh and Dr. Zhu-Liang Yang, Kunming Institute of Botany, Chinese Academy of
Sciences for reviewing of the manuscript. Thanks are also to Miss Li-Ming Wu who has
taken part in the collection and description of the new species.
Gloeocantharellus persicinus sp. nov. (China) ... 453
Literature cited
Ainsworth GC, Sparrow FK, Sussman AS. 1973. The fungi. An advanced treatise IV. B. Academic
Press. New York (USA). pp. 365-368.
Corner EJH. 1969. Notes on cantharelloid fungi. Nova Hedwigia 18: 783-818.
Kornerup A, Wanscher JH. 1978. Methuen handbook of colour. 3 ed. London: Methuen. 252 pp.
Lloyd CG. 1917. Mycological Notes, no. 47. Mycol. Writings 5: 653-668.
Pegler D. 1986. Agaric flora of Sri Lanka. Kew Bull. Addit. Ser. 12. pp. 455-456.
Petersen RH. 1971. The genera Gomphus and Gloeocantharellus in North America. Nova Hedwigia
21: 1-118.
_ Singer R. 1945. New genera of fungi-II. Lloydia 8: 139-144.
Smith AH, Hesler LR. 1943. New and interesting agarics from Tennessee and North Carolina.
Lloydia 6(4): 248-266.
Vasco-Palacios AM, Franco-Molano AE. 2005. A new species of Gloeocantharellus (Fungi-
Basidiomycetes) from Colombian Amazonia. Mycotaxon 91: 87-92
Yuan MS, Sun PQ. 1994. Sichuan mushrooms. Sichuan Science and Technology Press, Chengdu.
(in Chinese). p. 144, pl. 39.
Yuan MS, Sun PQ. 2007. Collection of coloured illustrations of Chinese mushrooms. Sichuan
Science and Technology Press, Chengdu. (in Chinese). p. 76, pl. 147.
MY COTA XON
Volume 106, pp. 455-467 October-December 2008
Critical review of recent records of
Russulaceae from Panama
F. HENNICKE & M. PIEPENBRING
florian. hennicke@uni-jena.de
Microbial Phytopathology, Institute of Microbiology
Friedrich Schiller-University of Jena
Neugasse 25, D-07743 Jena
piepenbr@daad-alumni.de
Department of Mycology, Institute of Ecology, Evolution and Diversity
Goethe-University of Frankfurt
Siesmayerstr. 70-72, D-60323 Frankfurt am Main
Abstract — Based on recently collected specimens, four new records of the Russulaceae
are presented for Panama: Russula aff. densissima, R. luteotacta, R. mexicana and
Lactarius aff. lilacinus. R. aff. densissima is reported for the first time for the Americas.
The Panamanian specimens of R. luteotacta, R. mexicana and L. aff. lilacinus are the
most southern records for the Americas. All species are described and illustrated. A
discussion of their known distribution, delimitation from morphologically similar
species, and their taxonomy is added, where applicable.
Key words — tropical mycology, Central America
Introduction
Tropical fungi are largely under-explored and the analysis of their diversity,
abundance, and taxonomy is far from complete. Especially taxonomically
challenging groups, like the Russulaceae, are badly known, because species
recognition and species demarcation demands a high degree of familiarity
with the group and the methods of identification. Only few monographs for
taxa outside Europe and North America are available (Buyck 1990, 1994, 1998,
Chiu 1945, Heim 1937, 1938a, b, 1955a, b, 1970, Hongo 1960, Karhula et al.
1998, Kong Luz 1995, McNabb 1973, Nuytinck et al. 2006a, b, Singer et al. 1983,
Verbeken 1996, 1998, 2001, Verbeken & Horak 2000, Verbeken et al. 2000).
These reasons make this group harder to approach than many other groups of
Basidiomycota.
Piepenbring (2006) assembled a checklist of fungi for Panama. With view
to the Russulaceae, she only listed four records of Russula spp. published by
Buyck & Ovrebo (2002). In order to identify recently collected specimens
456 ... Hennicke & Piepenbring
from Panama, European and American monographs of Russula (Bon 1988,
Einhellinger 1985, Romagnesi 1967, 1985, Sarnari 1998, 2005) and Lactarius
(Basso 1999, Heilmann-Claussen et al. 1998, Hesler & Smith 1979) as well as
fungus-floristic studies of other countries (Dennis 1970, Halling & Mueller
2005, Kimbrough 2000, Kreisel 1971, Mata 1999, Mata et al. 2003, Franco-
Molano et al. 2000, Pegler 1983, Singer 1975, Singer et al. 1983, Stevenson 1975)
including Russulaceae were used. In addition, a checklist of tropical Russula
spp. was consulted (Buyck 1992, 1993). The synonymy of the recorded species
is listed as far as the authors can agree to it.
Materials & methods
The present study is based on collections made by M. Piepenbring and coworkers
during their field work and mycological courses held at the Universidad
Autonoma de Chiriqui, in Panama. These specimens are deposited at the
National Herbarium of Panama (PMA) and the Botanische Staatssammlung
in Munich (M). They were analyzed macroscopically and microscopically with
a Carl Zeiss Stemi 2000 and a Jenalab microscope. Microphotographs were.
taken with a Sony 3 CCD-hig camara. SEM images were made at the Institute
for Systematic Zoology and Evolution at the FSU Jena. The identifications
were critically reviewed by R.-M. Rauschert (Freyburg, Germany), G. Weber
(Schwarmstedt, Germany) and A. Gminder (Jenapriefsnitz, Germany).
The Panamanian specimens were compared with specimens from personal
herbaria.
The known distribution of the identified species is presented based on
data from literature and specimen records deposited in the National Fungus
Collection of the U.S. Department of Agriculture (BPI), at the University of
Michigan Herbarium (MICH), the New York Botanical Garden (NY), and the
Oregon State University (OSC). These collections, however, were not analyzed
in the context of the present study.
Results & discussion
Lactarius
Lactarius aff. lilacinus (Lasch: Fr.) Fr., PLATE 1;,3: Fic. 1—5:; Fic
Epicr. syst. mycol. (Upsaliae): 348 (1838).
SYSTEMATIC POSITION (Heilmann-Claussen 1998): Subgenus Piperites, section Colorati
SPECIMEN VISUM: Panama: Chiriqui. Parque Nacional Volcén Baru, Sendero los
Quetzales, under Alnus acuminata Kunth, alt. approx. 2500 m, 20.X.2007, M. Piepenbring
with licenciatura students 4078 (PMA & M-0140998).
Macroscopic features: BASIDIOMATA (FIG. 1-2) grouped on soil under Alnus
acuminata; PiLeus 1.2-1.4 cm diam., brick to orange brown, surface dry
to somewhat viscid when wet, smooth, at first convex to applanate with a
decurved margin, later depressed, often remaining decurved, often with a small
Russulaceae of Panama... 457
umbo; STIPE 2.5-3 cm x 0.3-0.5 cm, cortex ochraceous to orange brown, with
a whitish rain-soaked tomentum-like not completely covering layer, beneath
it glabrous, overall cylindrical, often irregularly furrowed or rugged, base
rounded and dented; ConTeExtT fragile, becoming hollow in the stem, cream to
pale pinkish buff, in outer parts concolorous with the cortex, taste mild, slowly
becoming spicy to slightly bitter, smell not documented; latex rather sparse,
white; LAMELLAE pale pinkish buff, later apricot-ochraceous, broadly adnate
to decurrent, sometimes slightly intervenose, sometimes forked, crowded to
rather distant; SPORE PRINT not observed.
Microscopic features: SPoRES (Fic. 13) 5.9-8.7 x 5.1-6.9(-7.7) um, broadly
ellipsoidal to ellipsoidal, ornamentation more or less completely reticulate,
0.75-1 um high, of warts and short ridges in between, plage inamyloid; CysTIDIA
pleuromacrocystidia numerous, cylindrical, with an obtuse or somewhat
mucronate apex, cheilomacrocystidia (Fic. 5) rather numerous, cylindric
to slightly clavate, with an obtuse apex; PILEIPELLIS (Fic. 3-4) with poorly
developed cutis, oedotrichoderm with trichoderm-like parts, dermatocystidia
absent.
Hesler & Smith (1979) state the possibility of L. lilacinus occurring in North
America but they had never seen a specimen from North America when they
published their monograph. The herbarium (MICH), however, contains several
specimens of L. lilacinus collected in the eastern and the southeastern USA
from 1936 to 1939 (C.H. Kauffman. 19 Aug 1919, H.C. Beardslee Jr. 36069,
H.C. Beardslee Jr. 36137, H.C. Beardslee Jr. 39057). Most identifications made
by mycologists in the first half of last century — particularly those not familiar
with the origins of the fungal names they used — should be treated with
great skepticism. Thus, those USA records for L. lilacinus must be regarded as
doubtful. Lactarius lilacinus is well known from Europe (Heilmann-Claussen
et al. 1998, Basso 1999). To date, this species apparently has not yet been cited
for Central or South America. L. lilacinus is recorded here for the first time for
Central America.
L. lilacinus grows on damp soil in association with Alnus spp. Lactarius
species associated with Alnus may occur down to Colombia where Alnus has
its most Southern distribution in America.
In the past, the European L. lilacinus has been confused with L. spinosulus
Quél. because of the pinkish colors they both have. L. spinosulus is distinguished
by a stronger zonate and scalier cap. The species are easily separated by different
microscopic characteristics (Heilmann-Claussen et al. 1998). L. spinosulus has
not yet been documented for North America, neither in literature (Hesler &
Smith 1979) nor by the consulted herbaria. The pileus is somewhat orange in
the Panamanian specimen, which might cause confusion with other orange
species in the subgenus Russularia. Alder-associated species would include
L. lacunarum and the species around L. obscuratus — and possibly other species
458 ... Hennicke & Piepenbring
listed by Hesler & Smith (1979), Heilmann-Claussen et al. (1998) and Basso
(1999),
The Panamanian specimen, however, shows some deviating features
compared with literature data for L. lilacinus. Macroscopically the smaller
basidioma size is conspicuous when compared with the size as indicated in
literature, i.e. 1.5—8.0(-14.0) cm for the pileus and 1.5-5.5(-—7.0) x 0.3-2.0(-2.5)
cm for the stipe (Heilmann-Claussen et al. 1998, Basso 1999). This might be
explained by the tropical origin of the Panamanian specimen. ‘The availability
of mineral nutrients like soil-bound nitrogen, potassium, and phosphate often
limits the productivity of humid tropical forests (Lodge, 1993). Most nutrients
are concentrated in plant biomass and poor anion and cation exchange capacity
in the soils results in constant leaching out of the soils due to the high rainfall
(Jordan 1985, Lodge 1993). This is one possible explanation for smaller
basidioma sizes noted for tropical Russulaceae and other ectomycorrhizal
macrofungi strongly dependent on nutrition from their host trees.
Another deviation between the Panamanian specimen and typical L. lilacinus
collections is the absence of pinkish tinges in the pileipellis and the smoother,
hardly velutinous pileus surface (Heilmann-Claussen et al. 1998, Basso 1999).
Additionally, most basidiomata carry a small umbo upon their pileus. The
seemingly slightly shorter spores (Fic. 13) and the oedotrichoderm of the
Panamanian specimen contrast with literature descriptions for L. lilacinus. All
in all, these deviating characteristics create a habitus that is strongly reminiscent
of L. lacunarum Romagn. ex Hora but which is contradicted primarily by spore
measurements and the obtuse cheilomacrocystidia (Fic. 5). Latex colour changes
can be rather inconsistent, i.e. in the L. fulvissimus complex, a close relative of
L. lacunarum. Therefore we consider the observed differences in the latex colour
changeability as of minor importance. A bigger range of macromorphological
variation appears to be likely in tropical habitats because of the continuous
rainfalls and possibly because of the extremely heterogeneous conditions of
nutrient availability in the soil in tropical habitats. This makes microscopic data
more reliable. Thus, we identify the Panamanian specimen as L. aff. lilacinus.
More data from further specimens will show whether it corresponds to a new
forma or a new variety of this species.
Russula
Russula aff. densissima Jul. Schaff. ex Romagn., PLATE 1; 3: Fic. 6; Fic. 14-15
Bull. trimest. Soc. mycol. Fr. 96(3): 297 (1980).
Syn. Russula densifolia f. densissima Jul. Schaff. nom. inval.
? Russula fuliginosa Sarnari
SYSTEMATIC POSITION (Sarnari 1998): Subgenus Compactae, section Compactae, series
Acrifolia
SPECIMEN VISUM: Panama: Chiriqui. Parque Nacional Volcan Bart, Sendero los
Quetzales, alt. approx. 2500m, on soil under Quercus sp., 20.X.2007, M.Piepenbring
with licenciatura students 4085 (PMA & M-0140999),
Russulaceae of Panama... 459
PLaTE 1: 1-5 Lactarius aff. lilacinus (Piepenbring 4078). 1-2 Fresh specimen; 3 Longitudinal
section of pileipellis, scale 50 um; 4 Longitudinal section of pileipellis, terminal elements, scale
20 um; 5 Longitudinal section of lamellar edge, cheilomacrocystidia, scale 20 um; 6 Russula aff.
densissima (Piepenbring 4085). 6 Fresh specimen.
Macroscopic features: BASIDIOMATA (Fic. 6) solitary on soil under Quercus
sp.; PrLEUS 3 cm diam., brownish to pale grayish especially at the margin or
with yellowish to whitish specks, viscid and shiny when fresh, rapidly and
strongly blackening after injury or when old, depressed, later cyathiform,
margin rounded, later straightened; Stipe short, 2 x 1.3 cm, cortex white, then
sordid black or soot black blotchy, pruinose, slightly enlarged at the top, overall
nearly cylindrical, base rounded and dented; ConTExT white, with soot black
blotches in the stipe, at first becoming fuliginous, then blackening without
460 ... Hennicke & Piepenbring
reddening, firm and rigid, odor not distinct, taste acrid in the lamellae, mild
in the stipe; LAMELLAE whitish, becoming fuliginous and black in age or after
injury, decurrent or inserted obliquely, very rarely forked, very crowded (24-28
lamellae per cm); SPORE PRINT not obtained.
Microscopic features: SpoRES (Fic. 14) (6.2—)6.5-7.2(-9.2) x 5.6-6.5(-7.2)
um, subglobose, covered by an interrupted reticulum formed by low,
hemispherical warts 0.25-0.5 um high, partly interconnected; PILEIPELLIS
epicutis with cylindrical to clavate to subulate dermatocystidia (Fic. 15) with
hyaline or granular-refringent content in the segments.
Russula densissima has been described from France recently by Romagnesi
(1985). This species has only rarely been found by the authors of the European
Russula monographs. Romagnesi (1985) reported it from acidic soil under
different deciduous trees accompanied by some conifers, while Sarnari (1998)
collected it exclusively under Quercus sp. on clayey moderately calciferous soil.
The Panamanian specimen has also been found in association with Quercus sp.
Thus, it is likely that this species is neither strictly acidophilous nor dependant
on calciferous soil and it probably prefers Quercus spp. as mycorrhizal partner.
This is the first record of R. aff. densissima for the Americas, as there are no
records in literature or in the consulted herbaria. Central American records
of closely related species, such as Russula densifolia (Secr.) Gill., have been
reported by Gomez & Alfaro (1996) from Costa Rica. Other members of the
section Compactae ss. Sarnari resp. Nigricantae ss. Romagnesi (e.g. R. nigricans
(Bull.) Fr.) have also been found in North America. Corresponding specimens
have been deposited in the herbaria BPI, MICH, NY, and OSC.
R. densissima is difficult to distinguish from R. densifolia. In contrast
to R. densissima, R. densifolia has a reddening context, darker lamellae, and
less globose spores (Romagnesi, 1985). Discussing the variety of forms of
R. densifolia, Schaeffer (1952) already recognized hardly reddening rare
collections around Potsdam (Germany) with extremely crowded (16 lamellae
per 8 mm margin), slender, acutely marginate gills and only slightly acrid taste
with a strong smell of R. adusta Fr., which he preliminarily called R. densifolia
f. densissima. Another earlier monographic study of the section Compactae
of Russula (Shaffer 1962) primarily based on North American material also
analysed the critical group around R. densifolia. Discussing this issue, Shaffer
(1962) implies that the great variability of several features in all the different
forms of R. densifolia and adjacent species makes it difficult to base taxa only on
such differences. As an example several personal collections of R. densifolia with
mild taste and only slowly reddening basidiomata are mentioned that approach
R. adusta. The extremely crowded lamellae in R. densifolia f. densissima
compared with the variation of this feature in R. densifolia, for example, was
not considered significant by Shaffer (1962).
Russulaceae of Panama... 461
Sarnari (1998) suggests R. densissima as taxon affinum of R. fuliginosa but most
other specialists of Russula do not accept R. fuliginosa as a separate species
(G. Weber, pers. com.). R. densissima differs from R. fuliginosa in a smaller
sized pileus, subulate terminal segments of pileus hyphae (cylindrical in
R. fuliginosa), and the ~1-1.5 tm shorter spores. The fruiting body of the
specimen from Panama is smaller than in literature (Romagnesi 1985, Sarnari
1998), which should not be considered as unusual in view of its tropical montane
origin (comp. above). The spore measurements of the Panamanian specimen
are intermediate between R. densissima [6.5-7.2(-7.7) x 5.7-6.5(-7) um, cf.
Romagnesi 1985] and R. fuliginosa [7.2-8.8 x 5.6-7 um, cf. Sarnari 1998].
Russula luteotacta Rea, PLATE 2; 3: Fic. 7—9; Fic. 16-17
Brit. basidiomyc. (Cambridge): 469 (1922).
SYSTEMATIC POSITION (Sarnari 1998): Subgenus Russula, section Russula, subsection
Sardoninae, series Persicina
SPECIMEN VISUM: Panama: Chiriqui. Alto Chiquero, Finca de Maria Amoruso, alt.
approx. 1750 m, on soil under Quercus sp., 11.1X.2007, M. Piepenbring, J. Vega & A. del
Cid 3982 (PMA & M-0141000).
Macroscopic features: BASIDIOMATA (FIG. 7-9) grouped on soil under Quercus
sp.; PILEUS 4.5 cm diam., light red to pink, strongly discoloring by rain, dry,
shape first convex, later leveled and depressed in the center, margin rounded
and incurved; pileipellis strongly adhering, difficult to separate from the pileus
trama; STIPE 5.7 x 1.1 cm, club-shaped, cortex white, glabrous, base rounded,
at the beginning rigid, later spongy and fragile; CONTEXT white, not yellowing
when injured, spongy, odor not known, taste acrid; LAMELLAE (Fic. 9) white,
more or less decurrent, intervenose at the base, not forked, distant, not yellowing
when injured, fragile; SPORE PRINT white (Ia-b).
Microscopic features: SPORES (Fic. 16) (6.5-)7-8.5 (-9) x (5.5-)6-7 um,
mostly ellipsoidal, rarely subglobose, ornamentation verrucose to subspinose,
formed by mostly isolated elements, about 0.7 um high, connections rare and
short; PILEIPELLIS epicutis formed by dense, slender hyphae and cylindrical,
flexuous, very long and voluminous dermatocystidia (Fic. 17), dermatocystidia
obtuse or attenuated in an obtuse bottleneck, nearly constricted-capitate, often
septate, content of oily appearance, reacting greyish to sulfovanilline.
According to Singer et al. (1983), this species can also be found in the neotropical
lowlands. In Europe it is mostly associated with Carpinus spp., but can also be
found in association with other species of Fagales on clayey, sometimes soggy
soil (Romagnesi 1967, 1985, Singer et al. 1983). The Panamanian specimen is
the first record of Russula luteotacta for Panama and for Central and South
America, as no further American record than BPI 764448 from Maryland
(USA) could be found in literature or in herbaria.
462 ... Hennicke & Piepenbring
Taxonomists generally agree on the concept established for R. /uteotacta.
Lamellae mostly become chrome yellow when injured, but unchanging
collections are not rare (Sarnari 1998). The latter is also the case in the
Panamanian specimen - this is the only atypical feature of the collection. Other
features such as the distant lamellae, the color of the spore print, the fragility of
the fruiting body and the color of the pileipellis are typical.
Russula mexicana Butrl., PLATE 2; 3: Fig. 10—12; Fic. 18-19
Mycologia 3: 26 (1911). |
SYSTEMATIC POSITION (Sarnari 1998): Subgenus Russula, section Russula, subsection
Sardoninae, series Sanguinea
SPECIMEN VISUM: Panama: Chiriqui. Las Nubes, Alto Chiquero, close to the house of
ANAM, alt. approx. 1770 m, on soil under Quercus sp., 26. VIII.2007, M. Piepenbring,
with licenciatura students 3945 (PMA & M-0141001).
Macroscopic features: BASIDIOMATA (Frc. 10-11) grouped on soil under
Quercus sp.; PILEUS 4-6.5 cm diam., light red at the margin, center pallescent,
dry, shape first convex, than umbilicate, later more broadly depressed in the ~
center; STIPE 4-6.3 x 0.8-1.7 cm, cylindrical, cortex pinkish, glabrous, base
round; CONTEXT whitish, rigid, eventually becoming fragile in age, odor not
recognizable, taste acrid; LAMELLAE whitish, straightly to recessed adnate,
hardly intervenose, a few forked, close to moderately close; SPORE PRINT not
obtained.
Microscopic features: SPORES (FIG. 12, 19) 6.5-8 x 6-6.5 um, subglobose-
ellipsoidal, covered by blunt cylindrical to acute conical spines, mostly isolated
elements, 0.8-1.7 um high, scarcely interconnected by a few fine ridges;
PILEIPELLIS epicutis consisting of hyphal ends which in the central part of
the pileus mostly tend to be clavate, and of some dermatocystidia (Fic. 18)
with mostly rounded, subcapitate or slightly constricted terminal segments,
mostly with hyaline and sometimes granular content (terminal segment mostly
hyaline), reaction to sulfovanilline negative on desiccated material, staining
of acid resistant elements by carbolic fuchsine on the dry specimen showed a
granular incrustation by those elements at some of the dermatocystidia at the
subterminal and sometimes also at the terminal segments of the dermatocystidia
(Fic. 18).
The locality of the type (NY 653949) is situated in Mexico, near Jalapa. According
to Singer et al. (1983) this species seems to be common in mixed forests from
the Great Lakes region (e.g. Michigan, Illinois) south to subtropical and perhaps
tropical belt of North and Central America.
Singer et al. (1983) presumes that this species occurs in lowland pine and
oak forests south of Mexico, at least in plantations. The specimen Piepenbring
3945 is the first record of Russula mexicana for Panama and the most southern
Russulaceae of Panama... 463
PLATE 2: 7-9 Russula luteotacta (Piepenbring 3982). 7-9 Fresh specimen; 10-12 Russula mexicana
(Piepenbring 3945). 10-11 Fresh specimen; 12 SEM image of spores, scale 10 um.
report for this species in the Americas. Additional Central American records
are published by L. Lacey for Belize (http://web.cortland.edu/baronit/Belize/
list.asp) and Gémez & Alfaro (1996) for Costa Rica.
Several authors consider R. mexicana conspecific with R. sanguinea (Bull.)
Fr., as no significant differences could be found in the type specimens (Bills &
Miller 1984, Sarnari 1998), while other authors record them as separate species
(Singer et al. 1983, Gomez et al. 1996, G. Weber, pers. com.). Additionally,
Kong Luz (http://www.Russulamx.com) states that the type specimen of
R. mexicana is immature and therefore of limited use. The specimen from
464 ... Hennicke & Piepenbring
PLATE 3: 13 Lactarius aff. lilacinus (Piepenbring 4078). 13 Spore, scale 6,5 um; 14-15 Russula
aff. densissima (Piepenbring 4085). 14 Spore, scale 7 um; 15 Dermatocystidia, scale 20 um;
16-17 Russula luteotacta (Piepenbring 3982). 16 Spore, scale 6 um; 17 Dermatocystidia,
scale 10 um; 18-19 Russula mexicana (Piepenbring 3945). 18 Dermatocystidia, scale 10 um;
19 Spore, scale 6 um.
Panama corresponds to R. mexicana rather than R. sanguinea, chiefly
because of the shared habitat and apparent association with oaks. In contrast,
R. sanguinea grows exclusively under Pinus, Russula mexicana prefers montane
Quercus forests. Another striking difference is the spore ornamentation in
the Panamanian specimen, which consists of spines that are much longer and
rougher than in R. sanguinea, where the spines only reach up to 1 um in height.
Russulaceae of Panama ... 465
Apart from the spore ornamentation, the habitat is currently considered as the
only reliable feature for the separation of the two species.
Macroscopically R. mexicana differs by its glabrous stipe apex and its
thinner pileus cuticle (Singer et al. 1983). R. mexicana is supposed to have a
deep cream colored spore print of IId (Singer et al. 1983) while the spore print
of R. sanguinea is described as pale ochraceous (IIIa) (Bon 1988, Romagnesi
1985) or approximately IId (Sarnari 1998).
In general, the default assumption should be that the synonymy between
American and European taxa cannot be taken for granted, even in the absence of
clear morphological differences. The problem of a lack of significant differences
between the type of R. mexicana and R. sanguinea is certainly to be taken
seriously. Thus, it would be good to provide a modern, detailed description of
R. mexicana making this point. Such a study is in progress.
Acknowledgments
We would like to express our gratitude to Dr. U. Eberhardt (Utrecht, Netherlands) and
G. Weber (Schwarmstedt, Germany) for kindly reviewing the manuscript. We thank G.
Weber, R.-M. Rauschert and A. Gminder for helpful comments about the identifications
and the critical review of the descriptions. H. Dérfelt contributed photos of the specimens
and is thanked for critical reading of the manuscript. Assistance with REM imaging by
M. Eckart is acknowledged.
Fieldwork in Panama was successful with the help of students of Licenciatura of the
Universidad Auténoma de Chiriqui. The Autoridad Nacional del Ambiente (ANAM)
and M. Amoruso are thanked for permits to visit protected areas, the Herbario Nacional
de Panama (PMA) is thanked for logistic support.
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Vo
MYCOTAXON
Volume 106, pp. 469-477 October-December 2008
Three new species of Cortinarius subgenus Phlegmacium
TOBIAS GULDBERG FR@SLEV &
THOMAS STJERNEGAARD JEPPESEN
tobias. froeslev@gmail.com & thomas@stjernegaard. net
Botanical Museum
Gothersgade 130, DK-1123 Copenhagen K, Denmark
Abstract — Based on morphological and molecular phylogenetic studies we present
three new species in Cortinarius. Species descriptions are provided, along with
discussions of phylogenetic and morphological affinities to similar taxa. Cortinarius
majoranae, Cortinarius aquilanus and Cortinarius lepistoides spp. nov. are described.
The first taxon is placed in section Percomes, whereas the last two belong to section
Caerulescentes.
Keywords — taxonomy, phylogenetic relationships, European distribution
Introduction
Species delimitation of Cortinarius species is difficult, and the number of
accepted species and the application of names vary greatly in the modern
literature. Several recent studies of Cortinarius have employed phylogenetic
analyses of nuclear ribosomal gene sequence data from ITS (the internal
transcribed spacer region) and nLSU (nuclear large sub unit) in combination
with morphological studies with success (Ammirati et al. 2007, Froslev et al.
2005, 2006, 2007, Garnica et al. 2003a, b, 2005, Hoiland & Holst-Jensen 2000,
Kytovuori et al. 2005, Lindstr6m et al. 2008, Liu et al. 1997, Moser & Peintner
2002, Niskanen et al. 2006a, b, Peintner et al. 2001, 2003, 2004, Seidl 2000).
In this paper we present three new species of Cortinarius based on
morphological studies and phylogenetic analyses of the ITS region, Cortinarius
aquilanus, C. lepistoides and C. majoranae.
Material and methods
The taxonomic descriptions are primarily based on the material studied by the
authors. The measurements of macro morphological characters are based on
expanded, but never old (and then often aberrant) basidiocarps. Macro chemical
reagents applied were 2% and 40% KOH. The terminology of characters
follows Brandrud et al. (1990) with minor adjustments (see Brandrud 1996).
47() ... Froslev & Jeppesen
Microscopical structures were observed partly from fresh material mounted
in H,O, often with a drop of 40% KOH subsequently added, and partly from
dried material mounted in H,O and then KOH. The descriptions of the
pigment topography are based largely on observations from H,O mounts of
fresh, preferentially young material. The spores were studied and measured in
2 % KOH, with a 100x oil immersion lens. From each basidiocarp, a random
selection of mature spores obtained from cortina remnants were measured
excluding apiculus and ornamentation. Young, immature basidiocarps were
avoided. For species with many available specimens ten spores were. measured
per specimen. For species with few available collections thirty or more spores
were measured per specimen. Mean values (MV) of spore length and width as
well as Q—values (length/width ratio) were then calculated for each specimen.
MVs were calculated for each taxon. The pileipellis was studied from radial
(longitudinal), free hand sections, preferentially from fresh material. Wedge-
like sections, ultra thin at the one end, were obtained by cutting at a slightly
oblique angle. The sections were cut from approx. 5 x 5 x 3 mm large pieces
taken from young but expanded pilei, and at a position midway to the centre.
ITS sequence data obtained for this study were amplified and sequenced
according to the methods in Frgslev et al. (2005). Holotypes and representative
material of most relevant similar species were studied morphologically and
molecularly. Results of the molecular phylogenetic studies will not be presented
here. Collections marked with “tf” have been sequenced.
Herbarium acronyms follow Holmgren et al. (1990). The following acronyms
are used in collection numbers: TF=Tobias Guldberg Froslev and TSJ=Thomas
Stjernegaard Jeppesen. Colour photographs of most species and many of the
collections cited here are available on www.cortinarius.com (Fr@slev & Jeppesen
1999-2008).
Taxonomic descriptions
Cortinarius aquilanus T.S. Jeppesen & Froslev, sp. nov.
MycoBank MB 512354
Pileo 35-80 mm lato, hemisphaerico, dein plano-convexo, glutinoso, fibrilloso, ochraceo-
brunneo vel luteo-brunneo, ad marginem pallido ochraceo, KOH ope indistincto. Velo
universale sparso. Lamellis emarginatis, griseis. Stipite 35-65 x 8-20 mm, albido, dein
interdum luteo maculato, bulboso, bulbo distincte marginato, vel 35 mm lato. Carne albida,
in apice stipitis pallida grisea, sapore subnullo, KOH ope luteo in stipe, aurantio-brunneo
in bulbo, Sporis amygdaliformibus, ellipsoideis, distincte verrucosis, 9-10.5 x 5.5-6.5 um.
Typus: DENMARK: Horsens, Elbek skov, 21 Oct 2006, T.S.J., TSJ2006-110* (holotype C).
EryMo_coey: Latin - after eagle (Aquila), in reference to the locality, Ornereden (“The
eagle nest”) in Arhus, Denmark.
PILEus 3.5-8 cm, (hemi-)spherical, then plano-convex, glutinous, glabrous and
glossy, innately fibrillose, brown, ochre brown to yellow brown, margin pale
Cortinarius spp. nov. (Denmark & Sweden) ... 471
buff, slightly hygrophanous in very moist weather. Universal veil remnants fairly
sparse, and very thin with a “frosty” appearance (cf. Cortinarius glaucopus).
LAMELLAE crowded, emarginated, greyish white, becoming + ochre. STIPE
3.5-6.5 x 0.8-2.0 cm, with a sharply marginated bulb (-3.5 cm), whitish, with
age often staining somewhat yellowish in streaks or spots. CONTEXT whitish,
when young with + greyish tinge at stipe apex, becoming + yellow in damages.
SMELL + malty. Taste mild. MACROCHEMICAL REACTIONS: 2% and 40% KOH
on pileus negative to slightly brownish, in stipe context yellow to orange-yellow,
in bulb orange brown.
SPORES 9-10.5 x (5-)5.5-6.5 um (MV= 9.58 x 5.74 um, Q=1.67, n=61, 2
specimens), amygdaloid to ellipsoid, distinctly verrucose. PILEIPELLIS duplex:
cutis fairly thin, consisting of 10-15 layers of 2-4 um wide hyphae with zebra-
striped pigment incrustations. HyPODERM fairly weakly developed, of up to
12 um wide, sausage-shaped elements, embedded in yellow-brown parietal
pigment.
ECOLOGY AND DISTRIBUTION — Cortinarius aquilanus occurs in nemoral Fagus
sylvatica forests on calcareous or mineral rich soils.
SPECIMENS STUDIED — (*including macro characters): DENMARK: Fyn: Odense, Fruens
boge, 3 Oct 2000, T:S.J., TSJ2000-044*+ (herb C), Jutland: Arhusskovene, 3 Oct 2004,
T.S.J., TSJ2004-059*+ (herb C), loc. cit., 30 Aug 2005, T.S.J, TSJ2005-002*+ (herb C),
Horsens, Elbzek skov, 21 Oct 2006, T:S.J., TSJ2006-110*+ (holotype; herb C); Mon,
Klinteskov, 12 Sept 2006, T.F., TF2006-070*+ (herb. C). - SWEDEN: Skane: Degeberga,
HerremoOllan, 30. Sept 2003, T.S.J., TSJ2003-082* (herb. C)
COMMENTS —- Cortinarius aquilanus is a fairly anonymous Phlegmacium
species with pale stipe and lamellae and ochraceous brown to yellow brown
pileus, and a yellow to orange-yellow reaction in the context with KOH.
Cortinarius xanthoochraceus P.D. Orton is superficially very similar, but has
a more distinctly yellow, less fibrillose pileus, a negative reaction with KOH,
and smaller spores. Cortinarius multiformium Consiglio & Moénne-Locc.,
C. gracilior (M.M. Moser) M.M. Moser, and C. luteoimmarginatus Rob. Henry
differ by having citriform (papillate) spores. Cortinarius saporatus Britzelm. is
larger, has distinct velum remnants on a less fibrillose pileus, and larger, more
coarsely ornamented spores. Cortinarius langeorum Froslev & T.S. Jeppesen
is larger, has larger, more coarsely ornamented spores, and a reddish alkaline
reaction on the pileus. Cortinarius talus Fr. has a less distinct bulb, a honey-like
smell, and smaller spores. Cortinarius multiformis Fr. grows with conifers and
has a more or less honey-like smell.
The protologue of C. aurantiotinctus Bidaud in many ways corresponds
well with the present species and mentions a distinct KOH reaction. However,
the holotype of C. aurantiotinctus yielded an ITS sequence identical to
C. xanthoochraceus and should most likely be treated as a synonym hereof. We
472 ... Froslev & Jeppesen
do not know whether the protologue of C. aurantiotinctus is based on a mixed
concept or aberrant specimens of either species, but the holotype material
definitely does not belong with the present species.
Phylogenetically C. xanthoochraceus seems to be the closest relative of
C. aquilanus. The phylogenetic separation is unambiguous with more than
20 nucleotide differences and several indels. Cortinarius xanthoochraceus was
included in the study of Garnica et al. (2005) under the name C. langei (TUB
011861) and was placed close to the lineages Caerulescentes and Phlegmacioides.
A similar placement of both species is also indicated by our analyses.
The ITS sequence data were identical for all examined material. An ITS
sequence of the Danish holotype of C. aquilanus (TSJ2006-110) is deposited at
GenBank under the accession number FJ195642.
Cortinarius majoranae Froslev & TS. Jeppesen, sp. nov.
MycoBank MB 512355
Pileo 35-100 mm lato, hemisphaerico, dein plano-convexo, interdum umbonate, glutinoso,
primo luteo, dein ad centro olivaceo vel olivaceo-brunneo, minuto granulato, KOH ope
virido vel olivaceo. Velo universale indistincto. Lamellis emarginatis, viridi-luteis vel
luteis. Stipite 50-80 x 15-25 mm, clavato vel clavato-bulboso, bulbo interdum indistincto
marginate, virido-luteo vel Iuteo, ad basis e velo aurantio-luteo vel brueeno, mycelium
virido-luteo. Carne virida-lutea, sapore subnullo, KOH ope in stipite et pileo indistincto
olivaceo, in bulbo indistincto roseo. Odore distincto, ad C. percomis. Sporis ellipsoideis vel
amygdaliformibus, grosse verrucosis, 11-12.5 x 7-8 um. Typus: SWEDEN: Oland, Astad,
21 Sept 2003, T.S.J., TSJ2003-034* (holotype C).
ETyMoLoGy: Latin - after the marjoram-like smell
PILEus 5-10 cm, hemispherical to convex, then expanded, often with low umbo,
glutinous, glabrous and glossy, at first bright lemon yellow, soon becoming
olivaceous to olivaceous brown from centre, often +granulate-diffracted at
centre. LAMELLAE crowded, emarginate, greenish yellow to sulfur yellow.
STrPE 5-8 x 1.5-2.5 cm, clavate to + abruptly bulbous, greenish yellow to
sulfur yellow, in lower half with pale orange yellow to brownish veil remnants.
MYCELIAL STRANDS greenish yellow. CONTEXT greenish yellow. SMELL distinct,
like marjoram or apple-cake, similar to the smell of C. percomis, TAsTE mild but
with components of the smell. MACROCHEMICAL REACTIONS: KOH on pileus
greenish to olivaceous, in bulb context indistinctly reddish, in pileus and stipe
context indistinctly olivaceous.
SPORES 11-12.5 x (6.5-)7-8 um (MV=11.51 x 7.4 um, Q=1.55, n=45, two
specimens), ellipsoid to slightly amygdaloid, coarsely verrucose. PILEIPELLIS
simplex: Cutis thick, consisting of 30-35 hyphal layers. At surface 10-15
gelatinous layers of 2.5-3.5 um wide, loosely erect-entangled, hyaline hyphae.
The basal part of 20-30 layers of parallel to t+interwoven, 4—6 um wide hyphae.
Hyphae with intracellular granulate pigment, becoming wine red in 2% KOH.
:
|
Cortinarius spp. nov. (Denmark & Sweden) ... 473
ECOLOGY AND DISTRIBUTION — Cortinarius majoranae occurs in temperate
deciduous forests. It has been found associated with Tilia and Quercus on
calcareous soils, and seems to be extremely rare, but widespread. So far it has
been recorded in Sweden, the Czech Republic, and France.
SPECIMENS STUDIED — (*including macro characters): CZECH REPUBLIC: Prague,
Karlstejn, 21 Sept 2002, T.F & T.S.J., TSJ2002-018*+ (herb C). - SWEDEN: Oland,
Astad, 21 Sept 2003, T.S.J., TSJ2003-034*+ (holotype; herb. C). - FRANCE: Jura, Arbois,
3 Oct 1998, TE & T.S.J., TF1998-116 (herb. C), TSJ1998-135 (herb. C), loc. cit., 14 Oct
2000, T.S.J. & T.E, TF2000-098 (herb. C), TSJ2000-087 (herb. C), loc. cit., 1 Oct 2006,
T.SJ. & T.E, TSJ2006-083 (herb. C).
COMMENTS —- Cortinarius majoranae is easily recognized by the peculiar
(greenish) yellow basidiocarps and context, the clavate to slightly abruptly
bulbous stipe, and a smell similar to that of C. percomis. Cortinarius majoranae
has a greenish to olivaceous pileus and grows with frondose trees, whereas
C. percomis has an orange yellow pileus and grows exclusively with Picea.
Cortinarius aurilicis Chevassut & Trescol and C. xanthosuavis Bon & Trescol
occur in the same habitats as C. majoranae and can be very similar, but both
have a faint smell of banana peel, not of marjoram or apple-cake. Cortinarius
aurilicis furthermore has an umbra brown oxidation of the pileus and saffron-
orange mycelial strands. We have seen the species both with a clavate stipe
typical for species around C. percomis but also with an abruptly bulbous stipe.
The bulb is the considerably less developed than in species belonging to the
sections Caerulescentes/Glaucopodes and Calochroi/Fulvi, but comparable to
that of C. aurilicis.
The taxonomic status as a separate species was confirmed by a unique
phylogenetic position when comparing sequence data from C. majoranae with
representative specimens of most other putative taxa in the section Percomes.
Phylogenetically C. majoranae is placed in section Percomes along with
similar species such as C. nanceiensis Maire, C. aurilicis, C. percomis Fr.
and C. mussivus (Fr.) Melot, with which it shares the granulate diffracted
pileipellis, extractable, yellow, anthraquinonoid pigments, peculiar smells, and
a cylindrical to sub-bulbous stipe. It is phylogenetically distinct from all other
tested material from the section. It is placed as a sister to a lineage composed
of C. nanceiensis and C. aurilicis with more than 50 nucleotide differences to
any of these.
We did not succeed in amplifying ITS sequence data from the French
specimens. The Swedish and the Czech material were identical in the ITS
region. The ITS sequence data were identical for all examined material. An ITS
sequence of the Swedish holotype (TSJ2003-034) of C. majoranae is deposited
at GenBank under the accession number FJ195641. The holotype sequence
has partial ITS1 and ITS2 regions, and a more the more complete sequence of
the Czech collection (TSJ2002-018) is deposited under the accession number
FJ195640.
474 ... Froslev & Jeppesen
Cortinarius lepistoides TS. Jeppesen & Froslev, sp. nov.
MycoBank MB 512356
Pileo 30-70 mm lato, hemisphaerico, dein plano-convexo, glutinoso, primo violaceo-
caeruleo vel violaceo-griseo, dein griseo, ad centro brunneo-griseo, KOH ope luteo-
brunneo. Velo universale indistincto, brunneo. Lamellis emarginatis, griseis, ad marginem
distincto violaceo. Stipite 40-60 x 8-15 mm, caeruleo, dein pallido-caeruleo, bulboso,
bulbo distincte marginate, vel 25 mm lato. Margine bulbi e velo lilaceo-violaceo. Carne
albida, in apice stipitis caerulea, interdumin bulbo lutea, sapore subnullo, KOH ope luteo-
brunneo. Odore sub-nullo, Sporis limoniiformibus, distincte verrucosis, 8.5-10 x 5-6
um. Typus: DENMARK: Jylland: Vosnees Havskov, 30 Sept 2000, TF & TSJ, TF2000-056
(holotype C).
EryMo.oey: Latin, after Lepista, due to the superficial resemblance to Lepista nuda.
PrLEus 3-7 cm, hemispherical to convex, then expanded, glutinous, glabrous
and glossy, violaceous blue to dark violaceous grey, with radiate hygrophanous
streaks when young, soon becoming greyish with darker grey brown centre,
and + hygrophanous appearance, often with very thin and indistinct frosty veil,
becoming yellow brown under adhering grass, leaves etc. LAMELLAE greyish
with distinctly violaceous edge. STIPE 4-6 x 0.8-1.5 cm, with a marginate bulb
(-2.5 cm), blue at first, discolouring whitish from below, with violaceous blue
veil at bulb margin. CONTEXT whitish, in stipe bluish at first, soon becoming
whitish, becoming slightly yellowish in bulb. SMELL indistinct. TasTE mild.
MACROCHEMICAL REACTIONS: KOH on pileus yellow brown and in pileus
context, somewhat weaker in stipe and bulb.
SPORES 8.5-10 x 5-6 um (MV=9.38 x 5.48 um) Q=1.71 (n=90, two specimens),
limoniform, distinctly and densely verrucose. PILEIPELLIS duplex: Cutis fairly
thin, consisting of 10-15 layers of 2-4 um wide hyphae with intracellular
granulate pigment. HYPODERM well developed, consisting of up to 30 um wide,
subcellular elements, embedded in yellow-brown parietal pigment.
ECOLOGY AND DISTRIBUTION - Cortinarius lepistoides occurs in temperate
deciduous forests on calcareous soils. It is associated with Fagus sylvatica,
Quercus and possibly Corylus, and seems to be extremely rare but widespread
throughout Europe. It is known with certainty from Denmark, Sweden,
Germany and Hungary.
SPECIMENS STUDIED — (*including macro characters): DENMARK: Jutland, Vosnes
havskov, 17 Oct 1999, TF. & T.S.J., TSJ1999-094*+ (herb C), loc. cit., 30 Sept 2000, T.F.
& T.S.J., TF2000-056*+ (holotype; herb. C). - FRANCE: Dordogne, Notre-Dame-de-
Sanilhac, 4 Oct 2002, G. Eyssartier, GE02.074+ (herb. PC). - HUNGARY: Visegradi Mts,
Tahi, 11 Nov 2003, Balint Dima, DB628+ (herb. Dima). - SWEDEN: Oland, Odens, 3
Oct 1998, Pirjo & Ilkka Kytovuori, IK 98-2650 (herb. H).
COMMENTS - Cortinarius lepistoides is recognized by the initially blue pileus
becoming brownish and rust spotted, conspicuous violaceous lamellae edge,
and citriform spores. It is closely related to the similar species C. viridocoeruleus
Cortinarius spp. nov. (Denmark & Sweden) ... 475
Chevassut & Rob. Henry and C. camptoros Brandrud & Melot, but differs from
these by its initially violaceous blue pileus. Furthermore C. camptoros lacks
the distinctly violaceous lamellae edge seen in Cortinarius lepistoides and
C. viridocoeruleus. Cortinarius viridocoeruleus has larger spores than
C. lepistoides and C. camptoros.
Cortinarius imperialis Bidaudand C. subhygrophanus Bidaud showsimilarities
towards C. lepistoides, but the holotypes of both these species have amygdaloid
spores. Sequence data of C. subhygrophanus place this species very close to
Cortinarius arcifolius Rob. Henry, and also close to C. moénne-loccozii Bidaud,
but not as a relative of C. lepistoides. The holotype material of C. imperialis and
C. subhygrophanus originate from the same location, and the descriptions and
illustrations in Bidaud et al. (1993) indicate that they most likely represent the
same species, and therefore should be treated as synonyms.
Cortinarius lepistoides is little known, but has been included in some
publications. It was treated by Eyssartier (2004) under the name C. imperialis
(GE 02074) accompanied by a nice photograph and a discussion including
references to our initial publication of the species under the misapplied name
C. camptoros on www.cortinarius.com (Freslev & Jeppesen 1999-2008). An
examination of Eyssartier’s material revealed that it belongs to C. lepistoides.
The ITS data was identical to all other sequenced material and the spores were
limoniform and not amygdaloid as stated in the description in the text (Eyssartier
2004). An exact sequence match indicates that Cortinarius lepistoides was also
included in the study of Garnica et al. (2005) under the name C. viridocoeruleus
(TUB 011408, GenBank no# AY174788).
Phylogenetically it is most closely related to C. camptoros and C. virido-
coeruleus and placed in the phylogenetic lineage called Caerulescentes in Garnica
et al. (2005) containing species traditionally treated in sections Glaucopodes
and Caerulescentes. The species has three polymorphic sites in the ITS region of
which two show intragenomic polymorphisms as well, and the polymorphisms
do not divide the examined specimens into separate groups.
An ITS sequence of the Danish holotype (TF2000-056) of C. lepistoides is
deposited at GenBank under the accession number FJ195643. An ITS sequence
from a collection from the exact same locality (most likely the same mycelium)
the year before (TSJ1999-094) is deposited under the accession number
FJ195644. The two sequences are identical.
Acknowledgements
We thank B. Dima, G. Eyssartier, Ilkka Kytévuori, Kare Liimatainen, Tuula Niskanen
and J. Vesterholt for letting us study their collections and/or unpublished sequence
data, and the curators at C, G and P for arranging loans. Furthermore we thank Kare
Liimatainen and Tuula Niskanen for serving as pre-submission expert reviewers.
476 ... Froslev & Jeppesen
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of species in Cortinarius section Calochroi (Basidiomycota, Agaricales) in Europe. Molecular
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MYCOTAXON
Volume 106, pp. 479-483 October-December 2008
The first report of
Neoramularia bidentis from Europe
MAEGORZATA RUSZKIEWICZ-MICHALSKA' & AGATA WOECZANSKA2
*mrusz@biol.uni.lodz.pl
‘Department of Mycology, University of Lodz
Banacha 12/16, PL-90-237 Lodz, Poland
"Department of Botany and Mycology, Maria Curie-Sktodowska University
Akademicka 19, PL-20-033 Lublin, Poland
Abstract — Neoramularia bidentis, hitherto known only from a few Korean localities,
has recently been found in southern Poland. The habit, morphology and ecology of
this species and its host, Bidens tripartita, are described, discussed and compared with
Neoramularia spissa, and Ramularia concomitans, a similar hyphomycete on Bidens
species.
Key words — anamorphic fungi, hyphomycetes, distribution
Introduction
The genus Neoramularia U. Braun was introduced by Braun (1991). After the
modification of the circumscription and subsequent redispositions (Braun 1998,
Shin & Braun 1996), it now comprises ten species worldwide. They parasitize
mostly dicots, and only Neoramularia phragmitis (Nagorny) U. Braun has been
found on a member of the family Poaceae, viz. Phragmites australis (Cav.)
Trin ex Steud. (compare Table 1). While four Neoramularia species are known
exclusively from their type localities, Neoramularia kochiae (Woron.) U. Braun,
N. oxytropidis (Jacz.) U. Braun, N. phragmitis, N. rubi (Bubak) U. Braun and
N. spissa (Harkn.) U. Braun occur more abundantly (Braun 1998). N. bidentis
should now be added to this more widespread group.
The description of Neoramularia bidentis is based on material collected by
Shin in South Korea in 1992 (Shin & Braun 1993). Although it was repeatedly
observed in Kangung and Seoul in the following years (Shin & Kim 2001),
this species was not found outside South Korea (Farr et al. 2008). However,
Neoramularia bidentis has recently been collected in Poland during studies on
A480 ... Ruszkiewicz-Michalska & Wolczanska
micromycetes in southwest part of the country. It is the first confirmed locality
of this species both in Poland and in Europe.
Material and methods
Native to the Polish vascular plant flora (Mirek et al. 2002), Bidens tripartita
L. is widespread in the whole country (Zajac & Zajac 2001). This plant is a
permanent element of communities belonging to the Bidentetea class which
develop on drying shores of inland water bodies (Matuszkiewicz 2006). In
these habitats, Bidens tripartita coexists and competes with anthropophytes:
B. frondosa L., and, on a lesser scale, with B. connata Muhl. ex Willd. (Kucharski
1992), considered to be an alien invasive species (NOBANIS, www.nobanis.
org).
Infected host plants were collected in a poor meadow (Junco-Molinietum
association) with the participation of species of the Bidentetea class. The fungus
occurred abundantly on wild-growing Bidens tripartita from June to September
2006.
The collected host plants were air-dried and examined by standard light
microscopy (LM). Measurements of conidia and conidiophores, mounted in
lactophenol cotton blue and warmed, were taken. The monograph by Braun
(1998) was used for determination. The examined collections are deposited at
LOD and LBL.
TABLE 1. Host range and worldwide distribution of Neoramularia species*
NIASIE NACE NSE HOST (FAMILY) DISTRIBUTION
SPECIES
N. bidentis Bidens (Asteraceae) South Korea, Poland
N. capparis Capparis (Capparaceae) India
N. esfandiarii Scrophularia (Scrophulariaceae) Tran
N. karelii Vitex ( Verbenaceae) Turkey
N. kochiae Ceratocarpus, Eurotia, Kochia Central Asia, Caucasus, Europe
(Chenopodiaceae)
N. koreana Stachys (Lamiaceae) South Korea
N. oxytropidis Oxytropis (Fabaceae) Europe, North America
N. phragmitis Phragmites (Poaceae) Central Asia, Caucasus, Europe
N. rubi Rubus (Rosaceae) Central Asia, Caucasus, Europe
N. spissa Euthamia (Asteraceae) North America
*source: Braun (1998), Shin & Braun (1996).
Neoramularia bidentis new to Europe... 481
Fic. 1. Neoramularia bidentis: A, Leaf spots on Bidens tripartita; B, Conidiophores
and conidia; SCALE BAR = 20um; A. Wolczanska del.
Description and discussion
Neoramularia bidentis H.D. Shin & U. Braun, Mycotaxon 49: 352. 1993.
Leaf spots irregular, 2-15 x 2-5 mm, the largest one usually elongated, yellowish
brown with very thin, dark brownish margin (Fig. 1). Caespituli hypophyllous.
Conidiophores solitary or in small loose groups, erect, cylindrical with slightly
bulbose base, narrowed towards the tip, apex rounded, 18-32 x 4 um, up to 7
tum wide at the base (Braun 1998: 20-32(-55) x 4(-7) um). Conidiogenous loci
inconspicuous. Conidia 0-2-septate (mostly 1-septate), cylindrical or fusoid,
hyaline, 14-30 x 3.5-4.5(-—5) um (Braun 1998: 10-34 x (2.5-)3.5-5 um). Hila
neither thickened nor darkened.
SPECIMENS EXAMINED — POLAND. SLASKIE VOIVODESHIP: WOLA NEAR PSZCZYNA,
Junco-Molinietum association, on Bidens tripartita - 22.VI.2006; Ruszkiewicz-
Michalska & Myszka (LOD PF 2886); the same host, locality and collectors, 24. VII.2006,
28.VILI.2006, 10.[X.2006 (LOD PF 2913, 2934, 3081, LBL M 8652).
The Polish specimens agree well with the original description of N. bidentis
except for somewhat shorter conidiophores. Another Neoramularia species,
N. spissa, was also described on hosts of the family Asteraceae, but it clearly
differs from N. bidentis in having solitary, 2-celled conidia, 6-9(-—10) um wide.
Conidia of N. bidentis are catenate or solitary; they may be 1—3-celled and (2.5-)
3.5-5 um wide (Braun 1998). The distribution of these species is also different:
Neoramularia spissa is confined to North America (Canada, USA), while
N. bidentis is known from Asia (Korea) and has currently been found in Europe
(Poland).
482 ... Ruszkiewicz-Michalska & Wolczanska
In Poland, Ramularia concomitans Ellis & Holw. additionally occurs on Bidens
tripartita (Ruszkiewicz 2000, Wotczanska 2005). The morphology of hila and
conidiogenous loci, which are colorless and inconspicuous in Neoramularia and
conspicuously darkened in Ramularia, is the main feature that differentiates
the two genera.
The present collection of N. bidentis represents the first confirmed locality of this
species in Poland and in Europe at all. The specimen collected by Danilkiewicz
(1990) in Woroblin on the Bug River, published as Ramularia concomitans,
probably also belongs to this species (Wotczanska 2005).
Recent Polish reports of newly found phytopathogenic fungi mostly concern
taxa occurring either in plant communities barely researched by mycologists
or in urban areas. Our specimens of Neoramularia bidentis on Bidens
tripartita have also been recorded from habitats investigated less extensively
(Lawrynowicz et al. 2004). Only a few fungal species are known on this host
in Poland, namely Leptosphaeria ogilviensis (Berk. & Broome) Ces. & De Not.,
Septoria bidentis Sacc. (Mutenko & Majewski 2008), and a powdery mildew
reported by different authors as Sphaerotheca castagnei Lév., S. fuliginea
(Schltdl.) Pollacci, S. fusca (Fr.) Blumer emend. U. Braun as well as S. xanthii
(Castagne) L. Junell (Satata 1985, Mutenko 1989, Danilkiewicz 1990, Hotownia
& Kostrzewska 1991, Kalinowska-Kucharska & Kadtubowska 1993, Dynowska
et al. 1999, Czerniawska 2001). The herbarium material must be verified to
confirm the taxonomic status of these specimens in Podosphaera fusca (Fr.)
U. Braun & Shishkoff or P xanthii (Castagne) U. Braun & Shishkoff according
to the criteria proposed by Braun & Takamatsu (2000).
Acknowledgements
The authors are grateful to Prof. Dr. Uwe Braun (Halle/Saale, Germany) and to
Dr. Marcin Piatek (Krakow, Poland) for reading the manuscript and serving as pre-
submission reviewers. We are also indebted to Dr. Dorota Michalska-Hejduk (Lddz,
Poland) for phytosociological consultations.
The study was partially supported by the Polish Ministry of Science and Higher
Education (grant no 2 P04C 052 27).
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Volume 106, pp. 485-490 October-December 2008
Non-lichenized fungi from the Antarctic region
PAuL D. BRIDGE', BRIAN M. SPOONER? & PETER J. ROBERTS?
‘pdbr@bas.ac.uk
British Antarctic Survey, NERC, High Cross, Madingley Rd
Cambridge CB3 0ET, UK
*Mycology Section, Royal Botanic Gardens
Kew, Surrey TW9 3AB, UK
Abstract — A reference checklist has been compiled giving the names of all species
of non-lichenized fungi described by collection, observation or culture from the
general Antarctic region since records were first published in 1847. The list gives the
current names and taxonomic dispositions of over 1000 reported fungi (c. 750 species),
together with links to bibliographic references for each report. Additional data include
some details on the collection or isolation, and some limited information on associated
organisms and substrates. The list is maintained online via the Antarctic Environmental
Data Centre at: www.antarctica.ac.uk/bas_research/data/access/fungi/. The list is
updated on a regular basis.
Key words — continental Antarctica, sub-Antarctic, maritime Antarctica,
bibliography
Introduction
There is a long history of fungal recording from the Antarctic and sub-
Antarctic areas dating from the mid part of the 19" century. In 1847 Berkeley
described Peziza kerguelensis (= Scutellinia kerguelensis) as a new species from
Christmas Harbour, Kerguelen from material collected by Hooker in 1840
(Berkeley 1847). Berkeley later reported further material from Kerguelen that
had been collected during the HMS Challenger expedition of 1872-76 and by
Eaton during the Transit of Venus expedition of 1874-75 (Berkeley 1877a,b).
The first fungal material from the Antarctic mainland is generally accepted as
Sclerotium antarcticum (= Sclerotinia antarctica) collected in 1898 from Danco
Land on the Antarctic Peninsula during the expedition of SY Belgica (Bommer
& Rousseau 1905, Gamundi & Spinedi 1987).
There have now been over 200 publications describing or reporting fungi
from Antarctica since the early expeditions, and another 50-100 which are
otherwise relevant to mycological study of the region. Pegler et al. (1980)
486 ... Bridge, Spooner & Roberts
produced an annotated list of the macrofungi (Ascomycota and Basidiomycota)
described from the region up to 1979, including 14 new species based on
recently collected material. Some 90 species of largely filamentous microfungi
from terrestrial sites were listed by Vishniac (1996). Onofri et al. (2007) reported
251 species of fungi and oomycetes from mainland continental Antarctica,
but did not include details of species from the sub-Antarctic or many of the
islands between the sub- Antarctic and the mainland. To date there has been no
systematic listing of all of the 1000+ fungi reported from the region.
Methods and criteria
SCOPE AND souRCES. For this list the Antarctic is defined (following Pegler
et al. 1980) as the area south of 60°S, but also including Bouvet and the South
Sandwich Islands. The sub-Antarctic is taken as being the area between 60°S
and the Polar Front. The Polar Front is where warmer water from temperate
regions meets cold water from the Antarctic and gives rise to an upwelling. The
main information sources used were the libraries and herbaria of the British
Antarctic Survey (AAS) and the Royal Botanic Gardens, Kew (K(M)). The
main culture collections scanned were the American Type Culture Collection,
the Centraalbureau voor Schimmelcultures, and CABI Europe. Other sources
used included earlier unpublished literature searches and on-line resources
including the GBIF database (via www.gbif.org).
CONTENT AND ARRANGEMENT. Fungal data were recorded alphabetically in
single tables in html files. Full citations for the appropriate references were
included as an additional file. Each data entry was made up of the name of the
fungus as reported, its higher taxonomic position (e.g. order), an indication
as to whether the record was from the Antarctic or sub-Antarctic, its location,
and (where clearly evident) the substrate or associated organism. The reference
section provided details of at least one bibliographic reference that had
reported the taxon in the region. Papers describing subsequent work with the
collections outside the Antarctic region were not included to avoid duplication
of reports, and fossil fungi known from the region (see Onofri et al. 2007) are
also excluded. A small number of reviews were included, particularly where
they also included new observations or combinations (e.g. Pegler et al. 1980).
There are many additional publications relevant to the mycological study of
Antarctica and a compilation of these now under preparation will be added to
the web site in due course.
All taxa were initially listed under the names as reported. Species Fungorum
(www.indexfungorum.org/Names/Names.asp.) and the CBS yeast taxonomic
databases (www.cbs.knaw.nl/yeast/BioloMICS.aspx) were consulted to provide
current names, order level placements and synonyms. Where these have been
Antarctic fungi ... 487
considered synonyms or there was a later re-determination, the original entry
is limited to the name and original reference and hyperlinked to a full entry
under the current name. In a few instances where a name could not be traced
in the taxonomic literature or where there was some ambiguity in the original
report, a full data entry was included with a brief explanation.
DATA LIMITATIONS. Searches on the species epithets ‘antarctica’, “antarcticus
and ‘antarcticum’ were generally not useful as these epithets have been used
for collections from many southern hemisphere locations (e.g. Caretta &
Piotelli 1977). The most difficult data sets to interpret were those on the source/
substrate and the original location. In many instances sources were either not
clearly specified, or given as ‘soil’ with no further details. The detail regarding
location also varied considerably between reports, with some including specific
location information and others limited to the name of an island or a region
of the continent. In most cases this information is available in original or
associated publications and databases, and these categories in the list currently
largely contain minimum information that is gradually being improved at each
update.
Lichen-forming fungi have not been included in the list. Antarctic species
have been the subject of some contention (Dodge 1973, Castello & Nimis 1995,
Hawksworth & Iturriaga 2006), but they have recently been comprehensively
monographed by Ovstedal & Lewis Smith (2001, 2004) and a checklist has
been published by Olech (2001). An online checklist of Antarctic lichens is
also available at www.biologie.uni-hamburg.de/checklists/lichens/portalpages/
portalpage_checklists_switch.htm. Lichenicolous fungi are included here as
several reports and various names are not consistently listed in the above.
ACCESS AND USE. The list is available at the Antarctic Environmental Data
Centre, and can be accessed through the ‘Data and Collections’ page of the
main BAS internet site at www.antarctica.ac.uk/bas_research/data/index.php,
or directly at www.antarctica.ac.uk/bas_research/data/access/fungi/. The list
comprises an initial front page linked to further pages providing background
information and the data files. The list can be searched alphabetically or through
a separate index of genera. The full list of references included can be obtained
from the reference section of the supporting pages, or from the reference
heading on each data page. The list is reviewed and updated periodically as
data become available. The current version (2.3.1) is the sixth update since the
first posting in 2005.
Results
TAXA INCLUDED. Literature, herbarium and culture collection searches produced
a list of 1034 different fungal or oomycete and myxomycete names from 416
genera including 215 species with alternative current names in Index Fungorum.
488 ... Bridge, Spooner & Roberts
Of these, 884 had been identified at species level. All the major phyla within
the kingdom Fungi were represented, with the Ascomycota and Basidiomycota
the most frequent, accounting for 68% and 23% of the entries respectively.
The remainder belonged to the Zygomycota (5%) and Chytridiomycota (3%),
together with the non-fungal Myxomycota (1%) and Oomycota (1%). Some of
the species names listed were ambiguous or considered synonyms in one or
more reference source, and the final number of accepted species was estimated
at between 730 and 750.
BroGEoGRAPHY. Although the range and number of Antarctic fungal records
is relatively limited, there are some trends that can be identified from the data.
Larger fungi are mostly restricted to the sub-Antarctic and the islands. There
are a few reports of larger fungi on the mainland but most of these are restricted
to the Antarctic peninsular. At the current time the most southerly records
for larger fungi are long established colonies of two moss-associated Arrhenia/
Omphalina spp. on Leonie Island (67° 34’ S, 68° 08° W) currently under study.
Fungal diversity in the Antarctic appears significantly reduced in relation to
other regions. In very general terms individual niches appear to be occupied —
by only a few species, and the environmental factors defining the niche may be
more significant than the geography.
Eco.oey. The species that have been reported from the region include
representatives of most functional/ecological groups. Potential plant and
animal pathogens such as species of Pythium, Fusarium, Trichophyton,
Neozygites and Thyronectria species are present, as are groups normally
associated as mycorrhizae or plant or invertebrate symbionts. Relatively few
species have been reported directly from associated hosts, but examples include
Octospora leucoloma on moss covered soil, Neozygites sp. on mites, Paecilomyces
antarcticus on springtails and Pythium sp. on liverworts (see checklist). In
several instances species were not associated with their ‘usual’ function or
niche in the Antarctic environment. Examples include the normally insect-
associated species Lecanicillium lecanii that has been isolated from mosses
and gypsum crusts on rocks, the ericoid mycorrhizal species Rhizoscyphus
ericae that has been isolated from the roots of liverworts, and the normally
coprophilous Pirella circinans that has been found closely associated with beetle
cadavers on Bird Island (Hughes & Lawley 2003, Chambers et al. 1999, Bridge
et al. 2008). The mainly tropical and subtropical wood-rotting bracket fungus
Pycnoporus coccineus has been isolated from a mummified penguin. There have
been comparatively few fungal isolations from the marine environment, but
some of the more common genera identified from Antarctic sea water include
Aplanochytrium, Aniptodera, Camarosporium, Leucosporidium, Phoma and
Thraustochytrium (see checklist).
Antarctic fungi ... 489
Discussion
The condition ‘occurring in the Antarctic’ is here considered to include fungi
that are either clearly growing in the environment or are capable of surviving
at least transiently as spores or other propagules. As a result, the list is almost
entirely limited to recorded collections, observations or isolations. It does not
include data on fungi recorded solely by molecular methods, where there were
no other indications of their presence. Many of the fungal sequences obtained in
molecular diversity studies (e.g. Lawley et al. 2004) will undoubtedly come from
viable propagules or growing organisms, but interpretation and identification
can be problematic (see Bidartondo et al. 2008) so such data are generally not
included in the current version.
It is difficult to determine a precise level of endemism for Antarctic fungi.
Pegler et al. (1980) commented that the macrofungus biota of the sub- Antarctic
zone was remarkably similar to that of North Temperate- Arctic regions. Onofri
et al. (2007) listed some 22 species of microfungi that they considered endemic
to continental Antarctica, and a further 26 that are presumed able to grow there.
Some studies have looked at the potential transport of propagules, including
fungi, into the Antarctic in air movements. Chalmers et al. (1996) have shown
that air movements could transport fungi from South America to the Antarctic,
and that the meteorological conditions required would occur some three times
in every two years (Chalmers et al. 1996, Marshall 1996). Airborne distribution
would be less likely for many of the normally waterborne taxa such as Pythium
spp. De Hoog et al. (2005) have demonstrated that some species of Thelebolus
found in Antarctic lakes and deposits are also associated with birds, and such
vectors may be involved in the routine introduction of fungi. Given the wide
range of species reported in the region, and their generally cosmopolitan
distribution, it would seem likely that less than 2-3% of Antarctic fungi are truly
endemic, although some may occur in relatively long-established associations.
This is similar to the lower plants present where many have wide or bi-polar
distributions (Bednarek-Ochyra et al. 2000).
Acknowledgments
The authors thank J.A. Cooper and Paul M. Kirk for presubmission reviews.
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Cracow, Poland: Polish Academy of Sciences, Institute of Botany. 236 pp.
Berkeley MJ. 1847. Fungi. In Hooker JD. The Botany of the Antarctic Voyage of H.M. discovery
ships Erebus and Terror in the years 1839-43. Part 1(2): 447-454.
Berkeley MJ. 1877a. Enumeration of fungi collected during the expedition of H.M.S. Challenger.
(Second Notice). J. Linn. Soc., Bot. 15: 48-53.
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Berkeley MJ. 1877b. Report of the fungi collected in Kerguelen Island by the Rev. A.E. Eaton during
the stay of the transit-of-Venus expedition of 1874-75. J. Linn. Soc., Bot. 15: 221-222.
Bidartondo ML, et al. 2008. Preserving accuracy in GenBank. Science 319: 1616.
Bommer E, Rousseau M. 1905. Champignons. In Résultats du voyage du SY Belgica, expédition
Antarctique Belge 1897-1899. Rapports Scientifiques, Botanique. pp 1-5.
Bridge PD, Hughes KA, Denton JO. 2008. Association of the coprophilous fungus Pirella circinans
with an indigenous beetle on the sub-Antarctic Bird Island. Polar Biology DOI: 10.1007/
s00300-007-0403-y
Caretta G, Piotelli E. 1977. Microsporum magellanicum and Cunninghamella antarctica, new species
isolated from australic and Antarctic soil of Chile. Sabouraudia 15: 1-10. .
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Bibliotheca Lichenologica 57: 71-92.
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Antarctica. Mycological Research 103: 286-288.
de Hoog GS, Gottlich E, Platas G, Genilloud O, Leotta G, van Brummelen J. 2005. Evolution,
taxonomy and ecology of the genus Thelebolus in Antarctica. In Fungi of the Antarctic:
Evolution under extreme conditions (ed. GS de Hoog). Studies in Mycology 51: 33-76.
Dodge CW. 1973. Lichen Flora of the Antarctic Continent and Adjacent Islands. Phoenix Publishing:
Canaan, New Hampshire. 399pp
Gamundi JJ, Spinedi HA. 1987. Sclerotinia antarctica sp. nov., the teleomorph of the first fungus
described from Antarctica. Mycotaxon 29: 81-89.
Hawksworth DL, Iturriaga T. 2006. Lichenicolous fungi described from Antarctica and the sub-
Antarctic islands by Caroll W. Dodge (1895-1988). Antarctic Science 18: 291-301.
Hughes KA, Lawley B. 2003. A novel Antarctic microbial endolithic community within gypsum
crusts. Environmental Microbiology 5: 555-565.
Lawley B, Ripley S, Bridge P, Convey P. 2004. Molecular analysis of geographic patterns of eukaryotic
diversity in Antarctic soils. Applied & Environmental Microbiology 70: 5963-5972.
Marshall WA. 1996. Biological particles over Antarctica. Nature 383: 680.
Olech M. 2001. Annotated checklist of Antarctic lichens and lichenicolous fungi. Institute of
Botany of the Jagiellonian University: Krakow.
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Terra Antarctica Reports 11: 37-42.
Onofri, S, Zucconi L, Tosi S. 2007. Continental Antarctic fungi, [HW-Verlag
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identification and ecology. Cambridge University Press. 411pp
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South Georgia. Cryptogamie, Mycologie 25: 323-331.
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MYCOTAXON
Volume 106, pp. 491-501 October-December 2008
Villosiclava virens gen. nov., comb. nov.,
teleomorph of Ustilaginoidea virens,
the causal agent of rice false smut
Ey TANAKA’, TAKETO ASHIZAWA?’, RYOICHI SONODA; & CHIHIRO TANAKA*
tanakae@ishikawa-pu.ac.jp
'Department of Environmental Science e& Engineering, Ishikawa Prefectural Universit
P sy by y
Nonoichi-cho, Ishikawa 921-8836, Japan
Rice Disease Resistance Research Team, National Agricultural Research Center
Joetsu-shi, Niigata 943-0193, Japan
°Tea Pest Management Research Team, National Institute of Vegetable and Tea Science
Kanaya, Shimada-shi, Shizuoka 428-8501, Japan
*Graduate School of Agriculture, Kyoto University
Kitashirakawa Oiwake-cho, Sakyo-ku, Kyoto-shi 606-8502, Japan
Abstract—False smut is a serious worldwide rice disease. The anamorphic state of the
causal fungus is Ustilaginoidea virens. The teleomorphic state was introduced in 1934
and was allied with the family Clavicipitaceae. At present Claviceps virens is applied as the
teleomorph name of this fungus. However, morphological and biological characteristics
of the teleomorph are distinct from those of the genus Claviceps. In particular, the
Ustilaginoidea anamorph differs from the Sphacelia anamorph that characterises
the genus Claviceps. We deliberated on the nomenclature of rice false smut fungus,
compared the fungus with Claviceps, and found it is not appropriate to any existing
clavicipitaceous genera. In this paper, a new genus Villosiclava, a new combination
V. virens and an emended description for the teleomorph of rice false smut fungus are
proposed. In the absence of the original specimen, an epitype is also designated.
Key words—anamorph-teleomorph connection, Ascomycota, Hypocreales, mycotoxin,
taxonomy
Introduction
False smut is a serious disease in rice cultivation areas throughout the world.
The false smut disease affects the grains within the panicles and reduces rice
yield and quality (Ono & Uehara 1949). It has not attracted as much attention
as rice blast disease, partly because false smut and rice blast occur under
similar conditions in Japan (Yaegashi et al. 1989). In addition to affecting rice
production, water extracts of false smut diseased grains have been reported
492 ... Tanaka & al.
to poison animals (Nakamura et al. 1992, Suwa 1915). Furthermore, diseased
spikelets contaminate the rice harvest (Sonoda 1996).
The causal agent fungus forms so-called ‘green balls’ in place of a rice fruit.
Such balls are round or oblong in shape and uneven on their surface. The
pseudoscrelotia (green balls) consist of mycelial tissue and spore-masses and
incorporate remnants of anthers and portions of paleae and lemmas (Ikegami
1961). The spore masses are of orange to olive green colour, powdery and
somewhat sticky. The anamorph name of the causal fungus is Ustilaginoidea
virens. |
The teleomorph name of this fungus is currently Claviceps virens. However,
phylogenetic analyses have demonstrated that Ustilaginoideae species were not
allied with the genus Claviceps (Bischoff et al. 2004, Tanaka & Tanaka 2008,
White et al. 2000). Therefore, a taxonomic study on the teleomorph stage of this
fungus was performed and a new holomorphic name is here proposed.
Materials and methods
The ascigerous stage of rice false smut fungus was produced following previously
reported methods (Sakurai 1934, Hashioka et al. 1951) with slight modification.
True sclerotia were collected from rice spikelets on 8 October 2007 at Joetsu,
Niigata Prefecture, Japan (37° 15°38 °N, 138° 28° 58”E) by T. Ashizawa. The true
sclerotia were laid on moistened quartz-sand in glass petri dishes, maintained at
4°C for 1 month, and then placed at 24-28°C under fluorescent light and dark
conditions. Examined specimens were deposited in the mycological herbaria
of the National Museum of Nature and Science, Tokyo, Japan (TNS), and the
Natural History Museum and Institute, Chiba, Japan (CBM).
Vertical and longitudinal sections of mature capitula were obtained using
a paraffin embedding technique. Briefly, the fruiting bodies were fixed in FAA
solution [4% (v/v) formaldehyde, 5% (v/v) glacial acetic acid, and 50% (v/v)
ethanol], dehydrated, embedded in paraffin and sectioned at a 10 um thickness.
Microscopic characteristics were observed by differential interference contrast
microscopy.
Results and discussion
The ascigerous stage on the true sclerotia emerged during March to May 2008.
The characteristics of the fruiting bodies, ascostromata, perithecia, and asci
coincided with the descriptions of Sakurai (1934) and Hashioka (1971). The ~
true sclerotia generally had 1-5 stipitate, capitate fruiting bodies (Fic. 1A, B, C).
The fruiting bodies were initially bright yellow and then changed to dark green.
The stipes were shaggy and 5-15 mm in height (Fic. 1B, C, D). The ascostromata
were 1-3 mm in diameter. Perithecia were subcortical, and ovate to pyriform
Villosiclava virens gen. & comb. nov. (Japan) ... 493
Fic. 1. Teleomorph of Ustilaginoidea virens (TNS-F-18423). A: true sclerotia of various shapes;
B, C: stipitate, capitate ascostroma emerging from true sclerotia; D: ascostromata with shaggy
stipe; E, F: longitudinal sections of ascostromata; G: ascus with thickened apex; H, I: longitudinal
sections of perithecia ejecting ascospores through the ostiole; J, K: ascospores disarticulating into 4
part-spores and individual part-spores (arrows).
(Fic. 1E, EK H, I). Asci were numerous in a perithecium, hyaline, cylindrical,
with hemispherical and thickened apex, base attenuated and 8-spored (Fic.
1G, H, I). Ascospores were 140-230 um and disarticulated into 4 part-spores
which were hyaline, one-celled, filiform and 30-60 um x 1.3-1.8 um (ave. 45
x 1.5 um; Fie. 1J, K). Previous descriptions reported ascospores to be 120-180
494 ... Tanaka & al.
Fic. 2. Ustilaginoidea by Brefeld (1895), illustrated when the genus name Ustilaginoidea was
described. Pleurogenous thick-walled conidiospores and germination of thick-walled spores with
the formation of secondary conidiospores are shown.
um x 0.5-1.0 um (Sakurai 1934) or they were hyaline, one-celled, filiform and
50-80 um x 0.5-1.5 um (Hashioka 1971). These descriptions suggest that part-
spores were mistaken for ascospores. Thus, emendation of the description for
this fungus has been provided.
History of the rice false smut fungus
The anamorphic name of the rice false smut fungus is Ustilaginoidea virens.
Cooke (1878) first named the rice false smut fungus Ustilago virens, based on
‘a rice smut’ collected in India. Patouillard (1887) independently named the
fungus Tilletia oryzae, based on specimens collected in Japan. Brefeld (1895)
proved that Tilletia oryzae belonged to the fungi imperfecti, and not to the
basidiomycete family Ustilagineae. Brefeld (1895) proposed a new anamorphic
genus Ustilaginoidea (Fic. 2) based on the characteristics of the conidial stage
on rice and on Setaria crus-ardeae, and named the two species U. oryzae and
U. setariae, respectively. Takahashi (1896) pointed out that Ustilaginoidea oryzae
was identical with Ustilago virens, and thus combined the scientific name as
Ustilaginoidea virens, which was based on priority of epithet.
The teleomorph of the rice false smut fungus had not been found for
almost 40 years. Brefeld (1895) supposed that the rice false smut fungus was
an anamorphic state of Ascomycetes. Subsequently, Brefeld (1896) found
an ascigerous stage of U. setariae. He described, without illustrations, the
Villosiclava virens gen. & comb. nov. (Japan) ... 495
development of the fruiting bodies on true sclerotia and the fructification
of asci and ascospores. Brefeld (1896) also stated that the anamorphic genus
Ustilaginoidea was associated with Hypocreales. However, he did not propose a
new taxonomic name based on the teleomorph. Sakurai (1934) first found the
ascigerous stage of rice false smut fungus on true sclerotia. The true sclerotia,
which were hard, black and somewhat flattened, protruded from false smut
balls on rice. To date, the teleomorph of rice smut fungus has been reported in
Japan (Sakurai 1934, Hashioka et al. 1951), Korea (In et al. 1984), India (Singh
& Dubey 1984) and China (Wang et al. 1998). On the true sclerotia, stipitate
and capitate ascostromata develop (Sakurai 1934). Numerous perithecia
were embedded in the head of the fruiting body (Hashioka 1971). Sakurai
(1934) suggested that the rice false smut fungus was similar to Claviceps Tul.
(Clavicipitaceae), but did not propose a scientific name for the teleomorph.
Legitimacy of the current name of the rice false smut fungus
Claviceps oryzae-sativae (Hashioka 1971) has been often used as the
teleomorphic name of this rice false smut fungus (e.g. Bischoff et al. 2004,
Singh & Dubey 1984). However, Nakata (1934) already classified the fungus
as “Claviceps virens (Cke.) Sakurai” and provided a set of illustrations of the
teleomorph. While Nakata (1934) proposed the nomenclature only in Japanese,
the name fulfilled the requirements for valid publication at that time (McNeill
et al. 2006: ICBN Art. 36.1). This name is regarded as a formal error, because
the teleomorph is a new morph of the false smut fungus and such a morph
should be described as a new species (ICBN Art. 59.6). Therefore, this name
is attributed solely to Nakata (1934) and should be treated as Claviceps virens
M. Sakurai ex Nakata (ICBN Art. 46.4).
We recognized that the publication by Nakata (1934) was valid and that
Claviceps virens was the existing holomorphic name of the rice false smut fungus
being advocated. Nevertheless, Hashioka (1971) designated the teleomorph
of this fungus as Claviceps oryzae-sativae as an avowed substitute (nomen
novum), and treated the former Claviceps virens as a naked name (nomen
nudum). Hashioka (1971) did not indicate any reason for this treatment. We
suspect that he may have misapplied ICBN Art. 35.1 or misinterpreted Art. 36.1
when reviewing the publication by Nakata (1934). Alternatively, he may have
considered that Claviceps virens (Nakata 1934) was a provisional name and was
invalidly published (ICBN Art. 34), because Nakata (1937) in his second and
later editions of the same illustrated book classified the fungus as Ustilaginoidea
virens. However, because Nakata (1934) did not explicitly suggest that Claviceps
virens was a provisional name, it is not invalidated by ICBN Art. 34. In addition,
Hashioka (1971) did not designate a type, probably because “a new name
published as an avowed substitute for an older name is typified by the type of
496 ... Tanaka & al.
the older name” (ICBN Art. 7.3). Therefore, the name Claviceps oryzae-sativae
should be typified by the type of Claviceps virens. Thus the name Claviceps
oryzae-sativae is superfluous and therefore illegitimate (ICBN Art. 52.1).
The holotype was a set of illustrations drawn by M. Sakurai (Fic. 3) in the
publication by Nakata (1934). Unfortunately, the holotype illustration cannot
be critically identified for use in the precise application of the name of a taxon
(ICBN Art. 9.7). Thus, it is necessary to designate an epitype specimen in
support of the holotype.
Ahuja & Payak (1988) remarked that the false smut fungus should be
named Ustilaginoidea virens, not Claviceps oryzae-sativae. However, it was
inappropriate to apply the name Ustilaginoidea to a teleomorphic species at
that time, because the description of the genus Ustilaginoidea did not include a
teleomorph (Brefeld 1895). As a result, the genus Ustilaginoidea was applicable
only to the anamorph (ICBN Art. 59.3).
Taxonomic study
The rice false smut fungus differs from Claviceps purpurea, the type species of
the genus Claviceps, in several respects. The conidial state of most Claviceps
species is Sphacelia Lév., which is characterised by sporodochia with hard
hypostroma and honeydew-like exudates. The conidia are typically hyaline,
thin-walled and single-celled. The sclerotia of Claviceps, which give rise
to ascostromata, are irregularly globose, subglobose or elongated ovoid to
cylindrical in shape (Langdon 1954). In contrast, the rice false smut fungus
conidial state is Ustilaginoidea, which forms pleurogenous, sessile, pulverulent,
dark-coloured verrucose thick-walled conidiospores, and hyaline spherical
secondary conidiospores. The sclerotia are somewhat flat and irregular
oblong or horseshoe-shaped. In the Clavicipitaceae, teleomorph-anamorph
connections were thought important to their taxonomy (Diehl 1950), and that
suggestion was recently supported by molecular phylogeny (Kuldau et al. 1997,
Sung et al. 2007). Further molecular phylogenetic analyses have demonstrated
that Ustilaginoidea species are distinct from Claviceps (Bischoff et al. 2004,
Tanaka & Tanaka 2008, White et al. 2000). In addition, Claviceps species invade
host plants at the flowering stage, while the rice false smut fungus invades rice
plant at the booting stage or earlier (Fujita et al. 1989, Hisada 1936, Ikegami
1961). Claviceps species accumulate ergot alkaloids in the sclerotia; whereas,
the false smut fungus does not produce ergot alkaloids (Yamashita 1965) but-
produces several cyclic peptides, i.e., ustiloxins A-K which were isolated from
water extracts of false smut balls (Koiso et al. 1992, 1994, 1998; Li et al. 1995).
In the genus Claviceps, some poorly known species, such as C. flavella (Berk.
& M.A. Curtis) Petch, C. diadema (MOller) Diehl and C. orthocladae (Henn.)
Diehl, produce pseudosclerotia in grains (Pazoutova 2003, Diehl 1950). The
Villosiclava virens gen. & comb. nov. (Japan) ... 497
ced
=
3
iti 4
1. mE CR)
2. Fa
a. Hid
b, “FFE @ He AE
c. “RRO Br
d. FRE
e. -PRFRUT
CHIE)
psn oA
bs
Wee
Fic. 3. Illustration of teleomorph of rice false smut fungus by Sakurai as presented in Nakata (1934)
with permission from the publisher. Rice false smut disease; 1: diseased inflorescence (by Nakata).
2: causal fungus (by Sakurai). a: sclerotia; b: fruiting bodies; c: longitudinal section of ascostroma;
d: perithecia; e: ascus and ascospores (original legend in Japanese).
498 ... Tanaka & al.
mass of conidia in the host spikelet resembles the false smut ball of U. virens.
Further, C. orthocladae has yellow and villous stipitate ascostromata. However,
all these species have sphacelial anamorphs. Therefore, they are not congeneric
with the teleomorph of U. virens.
Thus, there exists numerous morphological, phylogenetic, ecological and
physiological differences between Claviceps and the teleomorph of U. virens.
There are no available genera possessing characteristics of the Ustilaginoidea
anamorph. For these reasons, we propose a new genus in the SPR to
accommodate holomorph of the rice false smut fungus.
Genus description
Villosiclava E. Tanaka & C. Tanaka, gen. nov.
MycoBank MB512265
SCLEROTIA applanata, inaequaliter oblonga. ASCOMATA capitata, stipitata, erecta e
sclerotio. Strp1TEs cylindrici, villosi. PERITHECIA ovato-pyriformia, partim vel omnino
in stromate immersa. Asci apice hemisphaerico et inspissato, cylindrici, hyalini, 8-spori.
Ascosporae filiformes, hyalinae. ANAMORPHE Ustilaginoidea.
SPECIES TYPICA: Villosiclava virens (Nakata) E. Tanaka & C. Tanaka
ETyMo oy: Latin, villosus, shaggy and clava, club; referring to Claviceps-like clavate
fruiting body with shaggy stipe.
SCLEROTIA flattened, irregular oblong. Ascomarta stipitate and capitate arising
from sclerotia. Stipes cylindrical and villose. PERITHECIA ovate-pyriform,
subcortical in ascomata. Asci cylindrical, hyaline, 8-spored, hemispherical at
the apex. Ascosporss filiform, hyaline. ANAMORPH Ustilaginoidea Bret. The
type species is Villosiclava virens.
CoMMENTS—The anamorphic state, Ustilaginoidea, is characterised by
pleurogenous pulverulent smut-like conidia occupying ovaries of Gramineae.
Primary conidia are globose to subglobose, yellow to dark-coloured and
verrucose. Secondary conidia are globose to ellipsoid, hyaline and smooth.
New combination and emended species description
Villosiclava virens (Nakata) E. Tanaka & C. Tanaka, comb. nov. etemend. (Fi. 1)
MycoBANnkK MB512266
Basionym: Claviceps virens M. Sakurai ex Nakata, Illustrations of Crop Diseases, Ist
Ed. 18 (1934)
= Claviceps oryzae-sativae Hashioka, Riso 20(3): 238 (1971) (nom. illegit.)
Anamorph: Ustilaginoidea virens (Cooke) Takah., Bot. Mag., Tokyo 10: 19 (1896)
= Ustilago virens Cooke, Grevillea 7: 15 (1878)
= Sphacelotheca virens (Cooke) Omori, Bot. Mag., Tokyo 10: 31 (1896)
= Tilletia oryzae Pat., Bull. Soc. Mycol. France 3: 119 (1887)
= Ustilaginoidea oryzae (Pat.) Bref., Unters. Gesammtgeb. Mykol. 12: 194-195 (1895)
= Ustilaginoidea albicans Shu Wang & J.K. Bai, Mycosystema 16: 257 (1997)
Species description by Nakata (1934) is emended as follows.
Villosiclava virens gen. & comb. noy. (Japan) ... 499
SCLEROTIA flat, botuliform, reniform, horseshoe-shaped, or differently shaped,
1 to several, usually 2, protruding from pseudosclerotia (false smut balls) on
spikelet of rice, usually 5-13 mm long, 2-5 mm wide, 1-2 mm thick. ASCOMATA
stipitate and capitate, 1-9 arising from the sclerotia. Stipes 5-15 mm long,
cylindrical, shaggy, phototropic, initially yellow then becoming dark-coloured.
CAPITULA globose, 1-3 mm in diameter, papillate when mature. PERITHECIA
ovate to pyriform, 150-350 um long, 70-150 um wide, embedded in the surface
of capitula, ostioles evidently erumpent. Asci cylindrical, hyaline, 130-300 um
long, 4-7 um wide with thickened apical cap. Ascospores filiform, hyaline,
eight per ascus, septate, 140-230 um long, 1.3-1.8 um wide, disarticulating at
septa to form 4 part-spores. PART-SPORES aseptate, 30-60 um long. ANAMORPH,
Ustilaginoidea virens.
SPECIMENS EXAMINED: TNS-F-18423—epitypus hic designatus, CBM-FA-37300,
originated from true sclerotia on grains of Oryza sativa, Joetsu-shi, Niigata, JAPAN.
Ex-epitype cultures originated from individual ascospores were deposited as MAFF
240994 and MAFF 240995 at the culture collections Genebank, National Institute of
Agrobiological Sciences, Tsukuba, Ibaraki, Japan.
The holotype supported by this epitypification is the illustration by Sakurai in
Nakata (1934) (q.v. Fic. 3). The original description in Japanese was published
in Nakata (1934). A Latin description was published in Hashioka (1971).
COMMENTS—At present, there is a single species in this genus. The teleomorph
of U. setariae is probably congeneric with V. virens, because it had true sclerotia,
yellow stipitate capitate ascostroma and ascospores disarticulating into 4 part-
spores (Brefeld 1896). Further studies are needed to obtain a conclusion. Other
fungi possessing the Ustilaginoidea anamorph may be included in this genus,
when an ascigerous stage is observed.
White mutants of rice false smut fungus have been occasionally reported
in Japan (Yaegashi et al. 1989, Honkura et al. 1991). Ustilaginoidea albicans,
reported from China (Wang & Bai 1997), is apparently a white mutant of the
rice false smut fungus. Both of these were characterised by their white colour
and the less verrucose surface of their spores (Honkura et al. 1991, Wang & Bai
1997).
The anamorphic genera Munkia Speg. and Neomunkia Petr. are
morphologically and phylogenetically close to Ustilaginoidea (Bischoff et al.
2004). Bischoff et al. (2004) suggested that the teleomorphic genus Mycomalus
Moller may be linked to Munkia martyris. Mycomalus, however, has no true
sclerotia and lacks stipitate ascostromata, but exhibits globose ascostromata on
bamboo stem, and their ascospores disarticulate into fusoid part-spores (Moller
1901). These characteristics are different from the teleomorph of U. virens.
500 ... Tanaka & al.
Acknowledgements
The authors thank Professor J. W. Spatafora and Dr. Y. Harada for reviewing the
manuscript. We also are indebted to Dr. S. Pennycook for very useful comments on
nomenclatural problems. We are grateful to Dr. H. Ikegami who kindly advised us on
some methods to produce the ascigerous stage of the rice false smut fungus.
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MYCOTAXON
Volume 106, pp. 503-504 October-December 2008
Validation of the basidiomycetous yeast species
Cryptococcus flavus and C. liquefaciens
ALVARO FONSECA", TEUN BOEKHOUT? & JACK W. FELL?
*amrf@fct.unl.pt
‘Centro de Recursos Microbiologicos (CREM)
Departamento de Ciéncias da Vida, Faculdade de Ciéncias e Tecnologia
Universidade Nova de Lisboa
Caparica, Portugal
?Centraalbureau voor Schimmelcultures
Utrecht, The Netherlands
Rosenstiel School of Marine and Atmospheric Sciences
University of Miami
Key Biscayne, Florida, U.S.A.
Abstract — Novel combinations are proposed to validate the binomials of two currently
accepted species belonging to the basidiomycetous yeast genus, Cryptococcus.
Key words — Basidiomycota, nomenclatural change
Introduction
When Phaff & Fell (1970) transferred Rhodotorula flava to the genus
Cryptococcus Vuill. as C. flavus, they failed to cite the basionym, Torula flava.
This omission rendered the new combination invalid (ICBN Art. 33.4; McNeil
et al. 2006). Nonetheless, the name was accepted in subsequent yeast taxonomy
treatises (e.g., Fell & Statzell-Tallman 1998, Rodrigues de Miranda 1984) and
was confirmed as a separate species by several molecular phylogenetic studies
(e.g., Fell et al. 2000, Scorzetti et al. 2002, Takashima & Nakase 1999).
Fonseca et al. (2000) transferred Torulopsis liquefaciens to the genus
Cryptococcus as C. liquefaciens, but an incorrect citation of the basionym
rendered the new combination invalid (ICBN Art. 33.4, Note 1). Cryptococcus
liquefaciens was confirmed as a separate species by subsequent molecular
phylogenetic studies (Scorzetti et al. 2002, Sugita et al. 2003). We hereby
propose two novel combinations to validate the binomials Cryptococcus flavus
and C. liquefaciens.
504 ... Fonseca, Boekhout & Fell
Nomenclatural changes
Cryptococcus flavus (Saito) A. Fonseca, Boekhout & Fell, comb. nov.
MycoBAank 512451
Basionym: Torula flava Saito, Japanese J. Botany 1(1): 45 (1922).
= Chromotorula flava (Saito) EC. Harrison (1928).
= Rhodotorula flava (Saito) Lodder (1934).
= Rhodotorula tokyoensis var. flava (Saito) T. Haseg. (1958).
= “Cryptococcus flavus” (Saito) Phaff & Fell (1970), nom. inval. (Art. 33.4).
Cryptococcus liquefaciens (Saito & M. Ota) A. Fonseca, Boekhout & Fell, comb. nov.
MycoBank 512452
Basionym: Torulopsis liquefaciens Saito & M. Ota, J. Brewery Science (Japan) 12: 167
(1934).
= “Cryptococcus liquefaciens” (Saito & M. Oda) A. Fonseca, Scorzetti & Fell (2000),
nom. inval. (Art. 33.4).
Acknowledgments
The authors thank Walter Gams and Cletus P. Kurtzman for presubmission review of the
manuscript and Shaun R. Pennycook for invaluable nomenclatural advice.
Literature cited
Fell JW, Statzell-Tallman A. 1998. Cryptococcus Vuillemin. pp. 742-767, in CP Kurtzman & JW Fell
(eds.), The Yeasts, a Taxonomic Study, 4th edn. Amsterdam, Elsevier.
Fell JW, Boekhout T, Fonseca A, Scorzetti G, Statzell-Tallman A. 2000. Biodiversity and systematics
of basidiomycetous yeasts as determined by large-subunit rDNA D1/D2 domain sequence
analysis. Int. J. Syst. Evol. Microbiol. 50: 1351-1371.
Fonseca A, Scorzetti G, Fell JW. 2000. Diversity in the yeast Cryptococcus albidus and related species
as revealed by ribosomal DNA sequence analysis. Can. J. Microbiol. 46: 7-27.
McNeill J, Barrie FE Burdet HM, Demoulin V, Hawksworth DL, Marhold K, Nicolson DH, Prado J,
Silva PC, Skog JE, Wiersema J, Turland NJ. 2006. International Code of Botanical Nomenclature
(Vienna Code). Adopted by the Seventeenth International Botanical Congress, Vienna, Austria,
July 2005. Regnum Vegetabile 146. 568 p.
Phaff HJ, Fell JW. 1970. Cryptococcus Kiitzing emend. Phaff et Spencer. pp. 1088-1145, in J Lodder
(ed.), The Yeasts, A Taxonomic Study, 2nd edn. Amsterdam, North-Holland.
Rodrigues de Miranda L. 1984. Cryptococcus Kiitzing emend. Phaff et Spencer. pp. 845-872, in
NJW Kreger-van Rij (ed.), The Yeasts, A Taxonomic Study, 3rd edn. Amsterdam, Elsevier.
Scorzetti G, Fell JW, Fonseca A, Statzell-Tallman A. 2002. Systematics of basidiomycetous yeasts:
a comparison of large subunit D1/D2 and internal transcribed spacer rDNA regions. FEMS
Yeast Res. 23495-5177;
Sugita T, Saito M, Ito T, Kato Y, Tsuboi R, Takeuchi S, Nishikawa A. 2003. The basidiomycetous ~
yeasts Cryptococcus diffluens and C. liquefaciens colonize the skin of patients with atopic
dermatitis. Microbiol. Immunol. 47: 945-950.
Takashima M, Nakase T. 1999. Molecular phylogeny of the genus Cryptococcus and related species
based on the sequences of SSU rDNA and internal transcribed spacer regions. Microbiol. Cult.
Coll. 15: 35-47.
505
NOMENCLATURAL NOVELTIES PROPOSED IN MYCOTAXON 106
Annulatascus apiculatus ER. Barbosa & Gusmao, p. 405
Bauerago tinantiae (J.C. Lindq.) Vanky, p. 162
Canoparmelia sanguinea Marcelli, Benatti & Elix, p. 436
Ceraceomyces bizonatus Dhingra & Avneet P. Singh, p. 399
Cercospora ochromae A. Hern.-Gut. & Dianese, p. 49
Cetraspora Oehl, EA. Souza & Sieverd., p. 337
Cetraspora armeniaca (Btaszk.) Oehl, EA. Souza & Sieverd., p. 338
Cetraspora gilmorei (Trappe & Gerd.) Oehl, FA. Souza & Sieverd., p.338
Cetraspora pellucida (T.H. Nicolson & N.C. Schenck) Oehl, F.A. Souza & Sieverd., p. 338
Cetraspora spinosissima (C. Walker & Cuenca) Oehl, F.A. Souza & Sieverd., p. 340
Cetraspora striata (Cuenca & R.A. Herrera). Oehl, FA. Souza & Sieverd., p. 340
Claviceps zizaniae (Fyles) Pantidou ex Redhead, M.E. Corlett & M.N.L. Lefebvre, p. 307
Cortinarius aquilanus T.S. Jeppesen & Fr@slev, p. 470
Cortinarius lepistoides T.S. Jeppesen & Froslev, p. 474
Cortinarius majoranae Froslev & T.S. Jeppesen, p. 472
Cryptococcus flavus (Saito) A. Fonseca, Boekhout & Fell, p. 504
Cryptococcus liquefaciens (Saito & M. Ota) A. Fonseca, Boekhout & Fell, p. 504
Dentiscutataceae F.A. Souza, Oehl & Sieverd., p. 340
Dentiscutata Sieverd., FA. Souza & Oehl, p. 340
Dentiscutata biornata (Spain, Sieverd. & S. Toro) Sieverd., FA. Souza & Oehl, p. 342
Dentiscutata cerradensis (Spain & J. Miranda) Sieverd., FA. Souza & Oehl, p. 342
Dentiscutata hawaiiensis (Koske & Gemma) Sieverd., RA. Souza & Oehl, p. 342
Dentiscutata heterogama (T.H. Nicolson & Gerd.) Sieverd., EA. Souza & Oehl, p. 342
Dentiscutata nigra (J.F. Redhead) Sieverd., EA. Souza & Oehl, p. 342
Dentiscutata reticulata (Koske, D.D. Mill. & C. Walker) Sieverd., EA. Souza & Oehl, p. 342
Dentiscutata scutata (C. Walker & Dieder.) Sieverd., FA. Souza & Oehl, p. 342
Dictyochaeta caatingensis A.C. Cruz & Gusmao, p. 16
Dictyochaeta intermedia Gusmao & S.M. Leao, p. 17
Dictyostelium culliculosum Yu Li & Xiao-Lan He, p. 380
Endocarpon crystallinum J.C. Wei & Jun Yang, p. 446
Entyloma arnaudianum (Cif.) Vanky, p. 158
Fuscutata Oehl, F.A. Souza & Sieverd., p. 342
Fuscutata heterogama Oehl, FE.A. Souza, L.C. Maia & Sieverd., p. 344
Fuscutata rubra (Stiirmer & J.B. Morton) Oehl, E.A. Souza & Sieverd., p. 347
Fuscutata savannicola (R.A. Herrera & Ferrer) Oehl, EA. Souza & Sieverd., p. 347
Fuscutata trirubiginopa (X.L. Pan & G.Yun Zhang) Oehl, FA. Souza & Sieverd., p. 347
506
Gjaerumia eremuri (Schwarzman) Vanky, 161
Gjaerumia muscari (Pass. ex J. Schrét.) Vanky, p. 161
Gloeocantharellus persicinus T.H. Li, Chun Y. Deng & L.M. Wu, p. 450
Glomus irregulare Blaszk., Wubet, Renker & Buscot, p. 252
Irpex hacksungii J.S. Lee & Y.W. Lim, p. 425
Kretzschmaria albogrisea Jad. Pereira, J.L. Bezerra & L.C. Maia, p. 238
Laetiporus montanus Cerny ex Tomiovsky & Jankovsky, p. 292
Leucoagaricus amazonicus A. Ortiz & Franco-Mol., p. 373
Ochrolechia pallentiisidiata Z.F. Jia & Q. Ren, p. 234
Paradoxa gigantospora (Y. Wang & Z.P. Li) Y. Wang, p. 200
Parmotrema sanctae-candidae Eliasaro, p. 432
Passalora schefflerae A. Hern.-Gut. & Dianese, p. 47
Pertusaria qilianensis Q. Ren & Z.T. Zhao, p. 441
Pertusaria paragilianensis Z.S. Sun & Z.T. Zhao, p. 443
Phaeoramularia cimicifugae FY. Zhai, Y.L. Guo & Yu Li, p. 203
Phallus luteus (Liou & L. Hwang) T. Kasuya, p. 8
Polyschema nigroseptatum R.M. Arias, R.F. Castaneda & Heredia, p. 30
Prathigada austroplenckiae A. Hern.-Gut. & Dianese, p. 57
Pseudocercospora astronii A. Hern.-Gut. & Dianese, p. 43
Pseudocercospora astroniiphila A. Hern.-Gut. & Dianese, p. 45
Pseudocercospora austroplenckiae A. Hern.-Gut. & Dianese, p. 59
Pseudocercospora eriothecae A. Hern.-Gut. & Dianese, p. 53
Pseudocercospora protii A. Hern.-Gut. & Dianese, p.55
Pseudocercospora pseudobombacis A. Hern.-Gut. & Dianese, p. 51
Quatunica F.A. Souza, Sieverd. & Oehl, p. 347
Quatunica erythropus (Koske & C. Walker) EA. Souza, Sieverd. & Oehl, p. 348
Racocetraceae Oehl, Sieverd. & F.A. Souza, p. 333
Racocetra Oehl, EA. Souza & Sieverd., p. 334
Racocetra alborosea (Ferrer & R.A. Herrera) Oehl, FA. Souza & Sieverd., p. 336
Racocetra castanea (C. Walker) Oehl, FA. Souza & Sieverd., p. 336
Racocetra coralloidea (Trappe, Gerd. & I. Ho) Oehl, FA. Souza & Sieverd., p. 336
Racocetra fulgida (Koske & C. Walker) Oehl, EA. Souza & Sieverd., p. 336
Racocetra gregaria (N.C. Schenck & T.H. Nicolson) Oehl, FA. Souza & Sieverd., p. 337.
Racocetra minuta (Ferrer & R.A. Herrera) Oehl, FA. Souza & Sieverd., p. 337
Racocetra persica (Koske & C. Walker) Oehl, FA. Souza & Sieverd., p. 337
Racocetra verrucosa (Koske & C. Walker) Oehl, EA. Souza & Sieverd., p. 337
Racocetra weresubiae (Koske & C. Walker) Oehl, FA. Souza & Sieverd., p. 337
507
Saccardoella psidiicola W.Y. Zhuang, W.Y. Li & K.D. Hyde, p. 414
Scutellosporaceae Sieverd., F.A. Souza & Oehl, p. 330
Scutellospora pernambucana Oehl, D.K. Silva, N. Freitas, L.C. Maia, p. 363
Sporisorium ciccaronei Vanky, p. 142
Sporisorium mandlaicum Vanky & N.D. Sharma, p. 141
Sporisorium shivasiorum Vanky, p. 145
Sporisorium stiparum (Speg.) Vanky, p. 163
Stereum hirsutum f. lobulatum Tura, Zmitr. & Wasser, p. 119
Thecaphora arnicae Vanky, p. 148
Thecaphora solidaginis (Ellis & Everh.) Vanky, p. 163
Tolyposporium isolepidis (Vanky) Vanky & M. Lutz, p. 155
Tranzscheliella distichlidis (McAlpine) Vanky, p. 137
Tranzscheliella iranica (Syd.) Vanky, p. 137
Tranzscheliella poae Vanky, p. 135
Tranzscheliella stipae-barbatae (Maire) Vanky, p. 138
Urocystis bulbinellae (P.H.B. Talbot) Vanky, M. Lutz, R. Bauer & Piatek, p. 154
Urocystis simplex (Liro) Vanky, p. 160
Ustilago milii-vernalis Vanky & H. Scholz, p. 148
Ustilago sclerachnes Wakef. ex Vanky, p. 150
Vanakripa rhizophorae R.M. Arias, Heredia & R.F. Castaneda, p. 34
Villosiclava E. Tanaka & C. Tanaka, p. 498
Villosiclava virens (Nakata) E. Tanaka & C. Tanaka, p. 498
Volvariella heterospora Menolli & Capelari, p. 389
Volvariella nullicystidiata Menolli & Capelari, p. 392
508
AUTHOR INDEX—VOLUME ONE HUNDRED SIX
Afshan, N.S., see Iqbal & al.
Arias Mota, Rosa Maria, Gabriela Heredia Abarca, Rafael F. Castaneda Ruiz & Cinthya
Ivonne Becerra Hernandez. Two new species of Polyschema and Vanakripa and
other microfungi recorded from mangrove in Veracruz, Mexico. 106: 29-40.
2008.
Ashizawa, Taketo, see Tanaka & al.
Bacci Jr., Mauricio, see Ortiz & al.
Barbosa, Flavia Rodrigues, Luis Fernando Pascholati Gusmao, Huzefa A. Raja &
Carol A. Shearer. Annulatascus apiculatus sp. nov., a new freshwater ascomycete
from the semi-arid Caatinga Biome of Brazil. 106: 403-407. 2008.
Barbosa, Flavia Rodrigues, see Cruz & al.
Baseia, Iuri Goulart & Francisco Diego Calonge. Calvatia sculpta, a striking puftball
occurring on Brazilian sand dunes. 106: 269-272. 2008. .
Baseia, [uri Goulart, see Cortez & al.
Baseia, I.G., see Oliveira & al.
Becerra Hernandez, Cinthya Ivonne, see Arias Mota & al.
Beltran-Tejera, E., see Telleria & al.
Benatti, Michel N., Marcelo P. Marcelli & John A. Elix. Canoparmelia sanguinea, a new
Parmeliaceae from Brazil. 106: 435-439. 2008.
Bezerra, José Luiz, see Pereira & al.
Blaszkowski, Janusz, Beata Czerniawska, Tesfaye Wubet, Tina Schafer, Francois Buscot
& Carsten Renker. Glomus irregulare, a new arbuscular mycorrhizal fungus in the
Glomeromycota. 106: 247-267. 2008.
Boekhout, Teun, see Fonseca & al.
Bridge, Paul D., Brian M. Spooner & Peter J. Roberts. Non-lichenized fungi from the
Antarctic region. 106: 485-490. 2008.
Buscot, Francois, see Blaszkowski & al.
Calonge, Francisco Diego, see Baseia & Calonge
Camino, M., G. Moreno, A. Castillo, D.W. Mitchell & D.W. Minter. Additions to the
myxomycete biota of Cuba. 1. 106: 75-102. 2008.
Campos-Santana, Marisa de & Clarice Loguercio-Leite. A note on Styptophyllum
erubescens. 106: 127-132. 2008.
Capelari, Marina, see Menolli & Capelari
Capelari, M., see Oliveira & al.
Cardoso, J., see Telleria & al.
Castaneda Ruiz, Rafael F., see Arias Mota & al.
Castillo, A. see Camino & al.
Castillo-Del-Moral, Ranulfo, see Guzman & al.
Cavalcanti, Maria A. Q., see Santiago & al.
Corlett, M.E., see Redhead & al.
—————
509
Cortez, Vagner Gularte, Marcelo Aloisio Sulzbacher, Iuri Goulart Baseia & Rosa Mara
Borges da Silveira. Two little known gasteroid fungi from Santa Catarina State,
southern Brazil. 106: 297-302. 2008.
Cruz, Alisson Cardoso Rodrigues da, Sheila Miranda Leao-Ferreira, Flavia Rodrigues
Barbosa & Luis Fernando Pascholati Gusmao. Conidial fungi from semi-arid
Caatinga biome of Brazil. New and interesting Dictyochaeta species. 106: 15-27.
2008.
Cuenca, Gisela, see Silva & al.
Czerniawska, Beata, see Blaszkowski & al.
Denchev, Cvetomir M., see Sesli & Denchev
Deng, Chun- Ying & Tai-Hui Li. Gloeocantharellus persicinus, a new species from China.
106: 449-453. 2008.
Deschamps, Jorge R., see Moreno & al.
de Souza, José Ivanildo, Iracema Helena Schoenlein-Crusius, Luisa Helena dos Santos
Oliveira, Norberto Carlos Schoenlein & Sandra Farto Botelho Trufem. Selected
species of Mucorales from soil contaminated with toxic metals in Sao Paulo State,
Brazil. 106: 273-288. 2008.
de Souza, Francisco Adriano, see Oehl & al.
Dhingra, G.S. & Avneet P. Singh. A new species of Ceraceomyces (Basidiomycetes) from
India. 106: 399-401. 2008.
Dianese, José C., see Hernandez-Gutiérrez & Dianese
Duenas, M., see Telleria & al.
Eliasaro, Sionara. Parmotrema sanctae-candidae, a new species in the Parmeliaceae
(lichenized Ascomycota) from Brazil. 106: 431-434. 2008.
Elix, John A., see Benatti & al.
Fell, Jack W., see Fonseca & al.
Fonseca, Alvaro, Teun Boekhout & Jack W. Fell. Validation of the basidiomycetous
yeast species Cryptococcus flavus and C. liquefaciens. 106: 503-504. 2008.
Franco-Molano, Ana Esperanza, see Ortiz & al.
Freitas, Nicacio de Oliveira, see Silva & al.
Froslev, Tobias Guldberg & Thomas Stjernegaard Jeppesen. Three new species of
Cortinarius subgenus Phlegmacium. 106: 469-477. 2008.
Gibertoni, Tatiana, see Martins-Junior & al.
Grand, L.F, C.S. Vernia & M.J. Munster. Biogeography and hosts of poroid wood
decay fungi in North Carolina: species of Trametes and Trichaptum. 106: 243-246.
2008.
Guo, Ying-Lan, see Zhai & al.
Gusmao, Luis Fernando Pascholati, see Barbosa & al.
Gusmao, Luis Fernando Pascholati, see Cruz & al.
Guzman, Gaston, Paul Kroeger, Florencia Ramirez-Guillén & Ranulfo Castillo-
Del-Moral. Species of Psilocybe in British Columbia, Canada (Basidiomycotina,
Agaricales, Strophariaceae). 106: 179-193. 2008.
Guzman, Gaston, see Medel & al.
510
He, Xiao-Lan & Yu Li. A new species of Dictyostelium. 106: 379-383. 2008.
Hennicke, F. & M. Piepenbring. Critical review of recent records of Russulaceae from
Panama. 106: 455-467. 2008.
Heredia Abarca, Gabriela, see Arias Mota & al.
Hernandez-Gutiérrez, Antonio & José C. Dianese. New cercosporoid fungi from the
Brazilian Cerrado 1. Species on hosts of the families Anacardiaceae, Araliaceae,
Bombacaceae, Burseraceae and Celastraceae. 106: 41-63. 2008.
Hu, Hung-Tao, see Wang & Hu
Illana, Carlos, see Moreno & al.
Iqbal, S.H. , A.N. Khalid, N.S. Afshan & A.R. Niazi. Rust fungi on Saccharum species
from Pakistan. 106: 219-226. 2008.
Jankovsky, Libor, see TomSovsky & Jankovsky
Jeppesen, Thomas Stjernegaard, see Froslev & Jeppesen
Jia, Ze-Feng, Qiang Ren & Zun-Tian Zhao. Ochrolechia pallentiisidiata, a new species
from China. 106: 233-236. 2008.
Kasuya, Taiga. Phallus luteus comb. nov., a new taxonomic treatment of a tropical
phalloid fungus. 106: 7-13. 2008.
Khalid, A.N., see Iqbal & al.
Kim, Changmu, see Lee & al.
Kroeger, Paul, see Guzman & al.
Lefebvre, M.N.L., see Redhead & al.
Leao-Ferreira, Sheila Miranda, see Cruz & al.
Lee, Jin Sung, Changmu Kim & Young Woon Lim. Irpex hacksungii sp. nov.
(Polyporaceae) from Korea. 106: 423-429. 2008.
Li, Tai-Hui, see Deng & Li
Li, Wen-Ying & Wen-Ying Zhuang. New species and new Chinese records of
Dothideomycetes. 106: 413-418. 2008.
Li, Yu, see He & Li
Li, Yu, see Zhai & al.
Lim, Young Woon, see Lee & al.
Liu, Ying-Jie, see Zhai & al.
Loguercio-Leite, Clarice, see Campos-Santana & Loguercio-Leite
Lorea-Hernandez, Francisco, see Medel & Lorea-Hernandez
Luo, Jing, see Zhuang & Luo
Maia, Leonor Costa, see Pereira & al.
Maia, Leonor Costa, see Silva & al.
Marcelli, Marcelo P., see Benatti & al.
Martins-Junior, Alcindo, Tatiana Gibertoni & Helen Sotao. Diplomitoporus
allantosporus (Basidiomycetes): a new record for Brazil. 106: 195-198. 2008.
Medel, Rosario & Francisco Lorea-Hernandez. Hyaloscyphaceae (Ascomycota) growing
on tree ferns in Mexico. 106: 209-217. 2008.
511
Medel, Rosario, Gaston Guzman & Florencia Ramirez-Guillén. First record of
Discoxylaria myrmecophila (Ascomycotina, Xylariales) from Veracruz, with new
reports from Jalisco, Morelos and Nuevo Leon (Mexico). 106: 1-6. 2008.
Melo, I., see Telleria & al.
Menolli Jr., Nelson & Marina Capelari. Records and two new species of Volvariella
(Pluteaceae, Agaricales) from Brazil. 106: 385-398. 2008.
Minter, D.W. see Camino & al.
Mitchell, D.W. see Camino & al.
Moreno, Gabriel, Carlos Illana & Jorge R. Deschamps. Comatricha argentinae, a
synonym of C. tenerrima. 106: 69-74. 2008.
Moreno, G., see Camino & al.
Munster, M.J., see Grand & al.
Nevo, Eviatar, see Tura & al.
Niazi, A.R., see Iqbal & al.
Oehl, Fritz, Francisco Adriano de Souza & Ewald Sieverding. Revision of Scutellospora
and description of five new genera and three new families in the arbuscular
mycorrhiza-forming Glomeromycetes. 106: 311-360. 2008.
Oehl, Fritz, see Silva & al.
Oliveira, J.J.S., C. Puccinelli, M. Capelari & I.G. Baseia. Neotypification of Marasmius
amazonicus. 106: 227-232. 2008.
Oliveira, Luisa Helena dos Santos, see de Souza & al.
Ortiz, Adriana, Ana Esperanza Franco-Molano & Mauricio Bacci Jr. A new species of
Leucoagaricus (Agaricaceae) from Colombia. 106: 371-378. 2008.
Pereira, Jadergudson, José Luiz Bezerra & Leonor Costa Maia. Kretzschmaria albogrisea
sp. nov. and K. curvirima from Brazil. 106: 237-241. 2008.
Piepenbring, M., see Hennicke & Piepenbring
Puccinelli, C., see Oliveira & al.
Raja, Huzefa A., see Barbosa & al.
Ramirez-Guillén, Florencia, see Guzman & al.
Ramirez-Guillén, Florencia, see Medel & al.
Redhead, S.A., M.E. Corlett, & M.N.L. Lefebvre. Validation and typification of the
name Claviceps zizaniae. 106: 303-309. 2008.
Ren, Qiang, Zhong-Shuai Sun & Zun-Tian Zhao. Two new species of Pertusaria
(Pertusariaceae) from China. 106: 441-444. 2008.
Ren, Qiang, see Jia & al.
Renker, Carsten, see Blaszkowski & al.
Roberts, Peter J., see Bridge & al.
Rodriguez-Armas, J.L., see Telleria & al.
Ruszkiewicz-Michalska, Malgorzata & Agata Wolczanska. The first report of
Neoramularia bidentis from Europe. 106: 479-483. 2008.
Salcedo, I., see Telleria & al.
ole
Santiago, André Luiz Cabral M. de A., Maria A. Q. Cavalcanti & Sandra Farto Botelho
Trufem. Mucor guilliermondii (Mucorales): a rare species found in herbivore dung
from Neotropics. 106: 103-108. 2008.
Schafer, Tina, see Blaszkowski & al.
Schoenlein, Norberto Carlos, see de Souza & al.
Schoenlein-Crusius, Iracema Helena, see de Souza & al.
Sesli, Ertugrul & Cvetomir M. Denchev. Checklists of the myxomycetes, larger
ascomycetes, and larger basidiomycetes in Turkey. 106: 65-67. 2008.
Shearer, Carol A., see Barbosa & al.
Sieverding, Ewald, see Oehl & al.
Silva, Danielle Karla Alves da, Nicacio de Oliveira Freitas, Gisela Cuenca, Leonor
Costa Maia & Fritz Oehl. Scutellospora pernambucana, a new fungal species in
the Glomeromycetes with a diagnostic germination orb. 106: 361-370. 2008.
Silveira, Rosa Mara Borges da, see Cortez & al.
Singh, Avneet P., see Dhingra & Singh
Sonoda, Ryoichi, see Tanaka & al.
Sotao, Helen, see Martins-Junior & al.
Spooner, Brian M., see Bridge & al.
Sulzbacher, Marcelo Aloisio, see Cortez & al.
Sun, Zhong-Shuai, see Ren & al.
Tanaka, Chihiro, see Tanaka & al.
Tanaka, Eiji, Taketo Ashizawa, Ryoichi Sonoda & Chihiro Tanaka. Villosiclava virens
gen. nov., comb. nov., teleomorph of Ustilaginoidea virens, the causal agent of rice
false smut. 106: 491-501. 2008.
Telleria, M.T., I. Melo, M. Duenias, I. Salcedo, J. Cardoso, J.L. Rodriguez-Armas &
E. Beltran-Tejera. Corticioid fungi (Basidiomycota) from Madeira Island. 106:
419-422. 2008.
TomSovsky, Michal & Libor Jankovsky. Validation and typification of Laetiporus
montanus. 106: 289-295. 2008.
Trufem, Sandra Farto Botelho, see de Souza & al.
Trufem, Sandra Farto Botelho, see Santiago & al.
Tura, Daniel, Ivan V. Zmitrovich, Solomon P. Wasser & Eviatar Nevo. The genus
Stereum in Israel. 106: 109-126. 2008.
Vanky, Kalman. Taxonomic studies on Ustilaginomycetes — 28. 106: 133-178. 2008.
Vernia, C.S., see Grand & al.
Wang, Yun & Hung-Tao Hu. Paradoxa gigantospora comb. nov. from China. 106:
199-202. 2008.
Wasser, Solomon P., see Tura & al.
Wei, Jiang-Chun, see Yang & Wei
Wolczanska, Agata, see Ruszkiewicz-Michalska & Wolczanska
Wubet, Tesfaye, see Blaszkowski & al.
ails:
Yang, Jun & Jiang-Chun Wei. Endocarpon crystallinum, the new lichen species from
semiarid deserts in China. 106: 445-448. 2008.
Zhai, Feng-Yan, Ying-Lan Guo, Ying-Jie Liu & Yu Li. A new species of Phaeoramularia
on Cimicifuga (Ranunculaceae). 106: 203-207. 2008.
Zhao, Zun-Tian, see Jia & al.
Zhao, Zun-Tian, see Ren & al.
Zhuang, Wen-Ying & Jing Luo. Re-identification of the anamorph of Leuconectria
grandis. 106: 409-412. 2008.
Zhuang, Wen- Ying, see Li & Zhuang
Zmitrovich, Ivan V., see Tura & al.
514
p.153, line 22
p.509, col. 1
front cover, legend
p.ili, line 1
p-194, lines 18-19
p-462, line 15
p.489, line 6
p.489, lines 27-28
p. 494, 3rd line up
ERRATA
VOLUME 85
for: micholsonii read: nicholsonii
for: Boubovia micholsonii read: Boubovia nicholsonii
VOLUME 105
for: Heterochaete tenuicula sp.nov. read: Heterochaete tenuicula (Lév.) Pat.
for: VOLUME ONE HUNDRED FIVE read: VOLUME ONE HUNDRED FOUR
for: Exsiccata 357, holotype; BPI 845215 & BPI 786230, isotype
read: Exsiccata 357 — BPI 845215 & BPI 786230, syntypes
for: Micarea melaena read: Micarea melaena
for: Arthonia hawksworthii Halici, p. 94
read: Arthonia hawksworthii Halici, p. 90
for: Leratiomyces similis (Pat. ex Sacc. & Trotter) Bresinsky & Manfr. Binder
ex Redhead & McNeill, p. 85
read: Leratiomyces similis (Pat. ex Sacc. & Trotter) Bresinsky & Manfr. Binder
ex Redhead & McNeill, p. 485
for: Takimichi Orihara read: Takamichi Orihara
back cover, 8th line up for: Takimichi Orihara read: Takamichi Orihara
ILD
REVIEWERS, VOLUME ONE HUNDRED SIX
The Editors express their appreciation to the following individuals who have, prior
to acceptance for publication, reviewed one or more of the papers prepared for this
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André Aptroot
Alan W. Archer
Richard E. Baird
Timothy J. Baroni
Gerald L. Benny
Annarosa Bernicchia
José Luis Bezerra
Uwe Braun
Marta Noemi Cabello
Lei Cai
Francisco de Diego
Calonge
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Sharon A. Cantrell
Lori M. Carris
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J.A. Cooper
Régis Courtecuisse
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Gisela Cuenca
Yu-Cheng Dai
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John A. Elix
Walter Gams
Aristoteles Gdes-Neto
Gaston Guzman
Laura Guzman-Davalos
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Roy E. Halling
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Richard C. Harris
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Hsiao-Man Ho
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Seppo Huhtinen
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Price aVickenzic
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Gabriel Moreno
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[Content continued from inside back cover]
Annulatascus apiculatus sp. nov., a new freshwater ascomycete from
the semi-arid Caatinga Biome of Brazil Flavia Rodrigues Barbosa,
| Luis Fernando Pascholati Gusmao, Huzefa A. Raja & Carol A. Shearer 403
Re-identification of the anamorph of Leuconectria grandis
Wen- Ying Zhuang & Jing Luo 409
New species and new Chinese records of Dothideomycetes
Wen- Ying Li & Wen-Ying Zhuang 413
Corticioid fungi (Basidiomycota) from Madeira Island
M.T. Telleria, I. Melo, M. Duenas. I. Salcedo,
J. Cardoso, J.L. Rodriguez-Armas & E. Beltran-Tejera 419
Irpex hacksungii sp. nov. (Polyporaceae) from Korea
Jin Sung Lee, Changmu Kim & Young Woon Lim 423
Parmotrema sanctae-candidae, a new species in the Parmeliaceae
(lichenized Ascomycota) from Brazil Sionara Eliasaro 431
Canoparmelia sanguinea, a new Parmeliaceae from Brazil
Michel N. Benatti, Marcelo P. Marcelli & John A. Elix 435
Two new species of Pertusaria (Pertusariaceae) from China
Qiang Ren, Zhong-Shuai Sun & Zun-Tian Zhao 441
Endocarpon crystallinum, the new lichen species from semiarid deserts
in China Jun Yang & Jiang-Chun Wei 445
Gloeocantharellus persicinus, a new species from China
Chun-Ying Deng & Tai-Hui Li 449
Critical review of recent records of Russulaceae from Panama
F. Hennicke & M. Piepenbring 455
Three new species of Cortinarius subgenus Phlegmacium
Tobias Guldberg Froslev & Thomas Stjernegaard Jeppesen 469
The first report of Neoramularia bidentis from Europe
Malgorzata Ruszkiewicz-Michalska & Agata Wolczanska 479
Non-lichenized fungi from the Antarctic region
Paul D. Bridge, Brian M. Spooner & Peter J. Roberts 485
Villosiclava virens gen. nov., comb. nov., the teleomorph of Ustilaginoidea virens,
the causal agent of rice false smut
Eiji Tanaka, Taketo Ashizawa, Ryoichi Sonoda & Chihiro Tanaka 491
Validation of the basidiomycetous yeast species Cryptococcus flavus and
C. liquefaciens Alvaro Fonseca, Teun Boekhout & Jack W. Fell 503
INDICES & INFORMATION
Nomenclatural novelties proposed in volume 106 505
Author index 508
Errata 514
Reviewers edhe
Submission procedures 516
[Content continued from back cover]
Neotypification of Marasmius amazonicus
J.J.S. Oliveira, C. Puccinelli, M. Capelari & I.G. Baseia 227
Ochrolechia pallentiisidiata, a new species from China
Ze-Feng Jia, Qiang Ren & Zun-Tian Zhao 233
Kretzschmaria albogrisea sp. nov. and K. curvirima from Brazil
Jadergudson Pereira, José Luiz Bezerra & Leonor Costa Maia 237
Biogeography and hosts of poroid wood decay fungi in North Carolina:
species of Trametes and Trichaptum
L.F. Grand, C.S. Vernia & M.J. Munster 243
Glomus irregulare, a new arbuscular mycorrhizal fungus in the
Glomeromycota Janusz Blaszkowski, Beata Czerniawska, Tesfaye Wubet,
Tina Schafer, Francois Buscot & Carsten Renker 247
Calvatia sculpta, a striking puffball occurring on Brazilian sand dunes
[uri Goulart Baseia & Francisco Diego Calonge 269
Selected species of Mucorales from soil contaminated with toxic metals
in Sao Paulo State, Brazil José Ivanildo de Souza,
Iracema Helena Schoenlein-Crusius, Luisa Helena dos Santos Oliveira,
Norberto Carlos Schoenlein & Sandra Farto Botelho Trufem 273
Validation and typification of Laetiporus montanus
Michal TomSovsky & Libor Jankovsky 289
Two little known gasteroid fungi from Santa Catarina State, southern Brazil
Vagner Gularte Cortez, Marcelo Aloisio Sulzbacher,
Iuri Goulart Baseia & Rosa Mara Borges da Silveira 297
Validation and typification of the name Claviceps zizaniae
S.A. Redhead, M.E. Corlett, & M.N.L. Lefebvre 303
Revision of Scutellospora and description of five new genera and three
new families in the arbuscular mycorrhiza-forming Glomeromycetes
Fritz Oehl, Francisco Adriano de Souza & Ewald Sieverding 311
Scutellospora pernambucana, a new fungal species in the Glomeromycetes
with a diagnostic germination orb Danielle Karla Alves da Silva,
Danielle Karla Alves da Silva, Nicacio de Oliveira Freitas,
Gisela Cuenca, Leonor Costa Maia & Fritz Oehl 361
A new species of Leucoagaricus (Agaricaceae) from Colombia
Adriana Ortiz, Ana Esperanza Franco-Molano & Mauricio Bacci Jr. 371
A new species of Dictyostelium Xiao-Lan He & Yu Li 379
Records and two new species of Volvariella (Pluteaceae, Agaricales)
from Brazil Nelson Menolli Jr. & Marina Capelari 385
A new species of Ceraceomyces (Basidiomycetes) from India
G.S. Dhingra & Avneet P. Singh 399
[Content continued opposite on page X]
“MYCOTAXON 106
SE “TABLEO OF CONTENTS
“First Serer of Discoxylaria myrmecophila (Ascomycotina, Xylariales) from
Veracruz, with new reports from Jalisco, Morelos and Nuevo Leon (Mexico)
aa RosarioMedel, Gast6n Guzman & Florencia Ramirez-Guillén
"Phallus luteus comb. nov., a new taxonomic treatment ay a tropical
» phalloid fungus. ERTL a tee ' Taiga Kasuya
- Conical fungi from semi- Said Caltinga biome of Brazil. New and interesting
_ Dictyochaeta species Alisson Cardoso Rodrigues da Cruz, Sheila Miranda
« Leao- Ferreira,Flavia Rodrigues Barbosa & Luis Fernando Pascholati Gusmao
“Two new species of Polyschema and Vanakripa. and other microfungi recorded
- from On eta in Veracruz, Mexico Rosa Maria Arias Mota, Gabriela Heredia
_ Abarca, Rafael F. Castafieda Ruiz & Cinthya Ivonne Becerra Hernandez
ses Hace fungi. from the Brazilian Cerrado 1. Species on hosts of
*, ae ithe families peeled oe Araliaceae, Bombacaceae, Burseraceae and
Tait -:Celastraceae Paes neg _ Antonio Hernandez- Gutiérrez & José C. Dianese
if if i ;Checkbas of ihe imyxomycetes larger ascomycetes, and larger basidiomycetes
He fii r “in ‘Turkey Beem ieran ee cerateeses 1 gud a Sesli & Cvetomir M. Denchev
_Comiatricha 1 argentinae,. a synonym of ee tenerrima a
abe - Gabriel Moreno, Carlos Illana & & Jorge R. Deschamps
us Additions to the ayect biota of Cuba. 1.
SRE EAL M. Camino, G. Moreno, A. Castillo, D. W. Mitchell & D.W. Minter
2 ae Macey pasliceniowl aMaaoraler) a rare species found in herbivore dung from
ra AeHOHIER es, ute André Luiz Cabral M. de A. Santiago,
1 faiientikey Breen, Chaat Be - Maria A. Q. Cavalcanti & Sandra Farto Botelho Trufem
“the e genus Stereum in Israel Ch SE Be Daniel Tura, Ivan V. Zmitrovich,
Gate ots _ Solomon Pp. hae & Eviatar Nevo
OM note ¢ on > Sopp duiberena: | : :
gest, Cee - Marisa de Campos- -Santana & Cine. Loguercio-Leite
isi taeonothi studies on i Dititagingricetes id EE Trees Kalman Vanky
: sons salves (Basidiomycotina, Agaricales, Strophariaceae) i in Canada, with a special
“review w of es from British Columbia Gast6n Guzman, Paul Kroeger,
| Florencia Ramirez- Guillén & Ranulfo Castillo-Del-Moral
~ Piplomitoporis either (Basidiomycetes): a new record for Brazil
_ Alcindo Martins-Junior, Tatiana Gibertoni & Helen Sotao 1°
Paani gigantospora comb. noy. from China Yun Wang & Hung-Tao Hu
Saat An new species of Phaeoramularia on Cimicifuga (Ranunculaceae)
Feng-Yan Zhai, Ying-Lan Guo, Ying- -Jie Liu & Yu Li
“Mpalsopase (Ascomycota) growing on treé ferns in Mexico
| : Rosario Medel & Francisco Lorea- Hernandez
Rust fungi o on. n Snecharum species from Pakistan
S.H. Iqbal, A.N. Khalid, N.S. Afshan & A.R. Niazi
[Content continues inside back cover] |
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